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UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 

AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  46 


Characters  Connected  With  the  Yield 
of  the  Corn  Plant 


(Publication  Authorized  August  6,  1921.) 


COLUMBIA,  MISSOURI 
AUGUST,  1921 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 

BOARD  OF  CONTROL 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY  P.  E.  BURTON  H.  J.  BLANTON 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 
OFFICERS  OF  THE  STATION 

J.  C.  JONES,  PH.  D.,  LL.  D.,  ACTING  PRESIDENT  OF  THE  UNIVERSITY 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 


STATION 

August, 


AGRICULTURAL  CHEMISTRY 
C.  R.  Moui/ton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  A.  M. 

E.  E.  Vanatta,  M.  S. 

R.  M.  Smith,  A.  M. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  SiEvEicing,  B.  S.  in  Agr. 

C.  F.  Ahmann,  A.  B. 

AGRICULTURAL  ENGINEERING 

J.  C.  Wooley,  B.  S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumeord,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

Paul  M.  Bernard,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

E.  F.  Hopkins,  Ph.  D. 

DAIRY  HUSBANDRY 

A.  C.  Ragsdale,  B.  S.  in  Agr. 

W.  W.  Swett,  A.  M. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride,  B.  S.  in  A. 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  A.  M. 

O.  W.  Letson,  A.  M. 

B.  B.  Branstetter,  B.  S.  in  Agr. 

B.  M.  King,  B.  S.  in  Agr. 

Alva  C.  Hill 


STAFF 

1921 

RURAL  LIFE 
O.  R.  Johnson,  A.  M. 

S.  D.  GromEr,  A.  M. 

Ben  H.  Frame,  B.  S.  in  Agr. 

FORESTRY 

Frederick  Dunlap,  F.  E. 

HORTICULTURE 

V.  R.  Gardner,  M.  S.  A. 

H.  D.  Hooker,  Jr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  M.  S. 

F.  C.  Bradford,  M.  S. 

H.  G.  Swartwout,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY 
H.  L.  Kempster,  B.  S. 

Earl  W.  Henderson 

SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M 

W.  A.  Albrecht,  Ph.  D. 

F.  L.  Duley,  A.  M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr 
Richard  Bradfield,  A.  B. 

O.  B.  Price,  B.  S.  in  Agr. 

veterinary  SCIENCE 
J.  W.  Connaway,  D.  V.  S.,  M.  D. 

L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 

ZOOLOGY 

George  Lefevre,  Ph.  D. 

OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Secretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Bertha  Hite,1  Seed  Analyst. 


Un  service  of  U.  S.  Department  of  Agriculture. 


CHARACTERS  CONNECTED  WITH  THE 
YIELD  OF  THE  CORN  PLANT 

W.  C.  Etheridge 


In  1909  the  Department  of  Agronomy  of  the  Missouri  Ex- 
periment Station  began  a study  of  the  factors  influencing  the  de- 
velopment of  the  corn  plant.  In  1914  this  department  was  divid- 
ed into  the  Departments  of  Soils  and  Field  Crops,  which  there- 
after separately  carried  on  those  phases  of  the  study  most  appro- 
priate for  their  respective  attention.  The  study  as  a whole  end- 
ed in  1920.  That  part  of  it  directly  concerned  with  the  effect 
of  nutrition  upon  growth  has  been  reported  by  Duley  and  Miller.1 
That  part  concerned  with  the  correlations  between  structure  and 
function  will  be  reported  in  this  paper.* 


I. — A Study  of  the  Correlation  Between  Yield 
and  Certain  Characters  of  the  Corn  Plant. 

The  essential  purpose  of  this  study  was  to  contribute  to  the 
solution  of  a problem  then  (1914)  receiving  much  attention  in 
the  field  of  plant  genetics — the  correlation  between  yield  and 
measurable  variations  in  the  visible  characters  of  the  corn  plant. 
It  is  a familiar  problem  to  all  who  have  read  closely  the  agro- 
nomic literature  of  the  past  12  years.  Likewise  its  conclusion, 
though  never  a real  solution,  is  nearly  conventional,  for  almost 
without  exception  its  investigators  have  reported  (1)  that  the 
correlations  did  not  exist  or  (2)  that  those  observed  were  not 
significant.  The  brief  results  reported  in  this  paper  are  not  ex- 
ceptional to  the  ensemble  of  evidence  from  similar  studies  by 
other  investigators.  They  are  reported  because  (1)  though  brief, 
they  contribute  a clear  case  and  (2)  the  great  weight  of  concor- 
dant evidence  now  existing  would  seem  almost  to  preclude  a pos- 


1This  and  subsequent  numerical  references  are  to  the  Bibliography. 

*The  writer  had  no  connection  with  this  project.  He  is  merely  a reporter  of  results  se- 
cured in  1910,  1911  and  1914,  from  studies  by  C.  B.  Hutchison,  C.  E.  Neff,  S.  B.  Nuck- 
ols  and  others.  His  presentations  and  interpretations  are  therefore  critical.  Possibly 
the  original  investigators  would  have  presented  their  data  more  accurately;  possibly  they 
would  have  interpreted  it  differently. 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  46 


sibility  that  further  study  of  the  problem  by  the  present  con- 
ventional methods  would  prove  fruitful. 

REVIEW  OF  RELATED  LITERATURE 

To  review  in  detail  the  evidence  contributed  by  previous  inves- 
tigators would  show  to  many  readers  a familiar  picture.  It  seems 
unnecessary  to  do  that.  Therefore  the  following  brief  summary 
presents  only  the  essential  developments. 

In  1909,  Montgomery2  reported  that  a long  (large)  ear,  medi 
um  depth  of  the  kernel  and  stockiness  of  the  stalk,  were  corre 
lated  with  relatively  high  yield.  Variation  in  other  characters  of 
the  plant  and  ear  showed  no  relation  to  yielding  ability.  The 
correlation  between  size  of  the  ear  and  yield  is  of  course  obvious 
— one  is  an  expression  of  the  other.  In  the  same  year  Hartley3 
reached  this  very  pointed  conclusion — “No  visible  characters  of 
apparently  good  seed  ears  are  indicative  of  high  yielding  power.” 
He  had  made  more  than  1,000  ear-rows  tests  of  4 varieties,  over 
a period  of  6 years. 

In  1910,  Pearl  and  Surface4  said  they  found  no  evidence  of 
close  association  between  the  conformation  of  the  seed  ear  and 
the  yield  that  it  produced.  They  had  studied  two  very  different 
types  of  sweet  corn,  giving  particular  attention  to  the  shape  of 
the  ear  and  the  covering  of  the  tip.  Ewing5  after  a very  thorough 
study  of  the  variation  in  several  dimensional  characters,  such  as 
height,  and  leaf  breadth,  concluded  that  “No  single  character 
among  those  studied  has  shown  itself  so  closely  connected  with 
yield  as  to  stand  out  as  a safe  guide  to  the  breeder.” 

In  1911,  Love6  concluded  that  no  characters  of  the  ear  were 
highly  correlated  with  earliness  and  that  none  could  serve  as  an 
index  of  earliness.  Sconce7  an  Illinois  seed  corn  breeder,  after  a 
study  of  6 years,  stated  his  belief  that  the  number  of  kernel-rows, 
the  form  of  the  kernel  and  the  size  of  the  germ  were  correlated 
with  the  yield  of  grain.  Funk8,  another  Illinois  seedsman,  while 
not  denying  the  existence  of  correlations,  concluded  that  the 
conventional  corn  score  card  does  not  emphasize  the  points  that 
affect  yield.  When  he  maintained  by  selection  the  type  which 
made  the  highest  yields,  he  gradually  produced  an  ear  very  diff- 
erent from  that  idealized  by  the  score  card. 

In  1913  McCall  and  Wheeler9  presented  their  interpretations 
of  various  statistical  data  of  other  investigators.  They  concluded 


Characters  Connected  with  the  Yield  of  the  Corn  Plant  5 


that  significant  correlations  between  yield  and  length,  weight, 
circumference  and  density  of  the  ear,  had  not  been  shown. 

In  1914  Williams  and  Welton10  made  an  exhaustive  report 
of  studies  through  a period  of  10  years.  As  an  average,  long  ears 
showed  an  advantage  in  acre  yield  of  1.39  bushels  over  short  ears; 
but  tapering  ears  showed  an  advantage  of  1.65  bushels  over  cylin- 
drical ears;  bare-tipped  ears  0.34  bushels  over  full-tipped  ears; 
smooth-dented  kernels  1.76  bushels  over  rough-dented  kernels; 
and  ears  of  a high  shelling  percentage  (88.16)  were  0.52  bushels 
lower  in  acre  yield  than  ears  with  a low  shelling  percentage  (76.07). 

In  1916,  Cunningham11  reported  that  smooth  and  medium 
smooth  kernels  outyielded  rough  kernels  by  a considerable  mar- 
gin. Variation  in  several  other  characters  showed  no  correla- 
tion with  yield.  He  concluded  that  correlations  were  variable  with 
the  environment. 

In  1917,  Cove  and  Wentz6  found  that  “The  characters  of 
length,  ratio  of  butt  to  tip,  average  circumference  of  cob,  weight 
of  ear,  average  weight  of  kernels,  number  of  rows  of  kernels,  and 
average  length  and  width  of  kernels  in  seed  ears  do  not  show 
correlations  significant  enough  to  be  of  value  in  judging  seed  corn.” 
They  reached  this  conclusion  after  five  years  of  study  with  one 
variety.  Hughes12  believed  the  first  year’s  results  of  his  experi- 
ment with  seed  corn  indicated  a close  correlation  between  yield 
and  the  ear  characters  idealized  by  the  score  card. 

In  1918  Hutcheson  and  Wolfe13  believed  they  had  found  sig- 
nificant correlations  between  yield  and  the  size  and  trueness  to 
type  of  the  ears.  Many  other  characters,  such  as  shelling  per- 
centage, number  of  rows,  space  between  rows,  and  the  filling  of 
the  butt,  were  not  related  to  yield  in  a significant  way.  Olson, 
Bull  and  Hayes14  from  apparently  the  soundest  and  most  com- 
prehensive study  yet  conducted,  failed  to  find  a significant  cor- 
relation between  yield  and  any  of  a broad  range  of  characters  ob- 
served. They  made  the  very  practical  statement  that  “Close  se- 
lection for  high  scoring  ears  is  of  no  practical  value  in  increas- 
ing  yield.” 


6 Missouri  Agr.  Exp.  Sta.  Research  Bulletin  46 

MATERIAL  AND  METHODS 

Ten  ears  of  each  type  in  the  following  groups  were  selected 
from  the  variety  Boone  County  White,  as  seed  for  the  crop  in 
which  the  correlation  study  was  to  be  made. 

A.  Long,  slim,  smooth  ears 

B.  Long,  thick,  smooth  ears 

C.  Short,  thick,  smooth  ears 

D.  Medium  long,  medium  thick,  medium  rough 

E.  Long,  slim,  rough  ears 

F.  Long,  thick,  rough  ears 

G.  Short,  thick,  rough  ears 

Each  of  the  70  ears  was  planted  in  an  ear-row,  each  type  in  a 
separate  series.  Thus  there  were  10  ear-rows  of  long,  slim,  smooth 
ears ; 10  of  long,  thick,  smooth  ears ; and  so  on.  The  series  were 
contiguous  and  each  fifth  row  was  a check,  planted  with  seed  of 
one  type.  The  hills  were  spaced  44  inches  apart,  each  way,  and 
two  plants  -were  left  in  each  hill  as  the  final  stand.  The  crop  re- 
ceived ordinary  cultivation.  Just  before  the  tasseling  stage  40 
normal  plants  in  each  row  were  labeled,  each  plant  standing  among 
similar  normal  plants.  There  were  labeled  a total  of  2800  plants 
and  for  each  the  following  data  were  recorded : 

1.  Date  of  tasseling. 

2.  Date  of  silking. 

3.  Number  of  tillers  at  full  growth. 

4.  Leaf  area  above  the  ear  at  full  growth. 

5.  Leaf  area  below  the  ear  at  full  growth. 

6.  Total  leaf  area  at  full  growth. 

7.  Height  of  ear  at  full  growth. 

8.  Relative  position  of  the  ear-shank. 

9.  Height  of  stalk  at  full  growth. 

10.  Number  of  nodes  in  stalk. 

11.  Circumference  of  first  internode  above  ground,  at 
full  growth. 

12.  Circumference  of  first  internode  above  ear,  at  full 
growth. 

13.  Tassel  length. 

When  two  ears  were  borne  by  a plant,  all  measurements  with 
reference  to  the  ear  were  made  by  the  upper  ear  only,  although  the 
total  yield  of  both  ears  was  determined.  The  leaf  area  was  the 


Characters  Connected  with  the  Yield  of  the  Corn  Plant  7 


sum  of  the  areas  of  individual  leaves  measured  by  Montgomery’s 
formula — Area=12x^4  (breadth  x length).  The  tassel  length  was 
measured  by  finding  the  sum  of  the  length  of  five  average  lateral 
branches,  dividing  this  sum  by  five  and  multiplying  the  quotient  by 
the  number  of  laterals,  then  adding  to  the  product  the  length  of  the 
central  spike.  Sound  ears  were  gathered  from  1,761  of  the  2,800 
plants  measured,  and  stored  under  good  drying  conditions  for 
six  weeks.  The  weight  of  shelled  grain  produced  by  each  plant 
was  computed  on  the  basis  of  a uniform  content  of  moisture. 

THE  RESULTS 

The  mean  yields  of  shelled  grain  produced  by  plants  from 
the  various  types  of  seed-ears  are  shown  here. 


Table  1. — The  Relative  Productivity  oe  Seed  From  DieeerEnt  Types  oe  Ears. 


Series  Type  of  original 
seed  ear 

No.  of  ears 
harvested  for 
yield  test 

Mean  yield  in 
ounces  of  shelled 
grain  per  plant 

Probable 

error 

(±) 

A 

Long,  slim,  smooth 

195 

7.7400 

.1409 

B 

Long,  thick,  smooth 

228 

7.7150 

.1169 

C 

Short,  thick,  smooth 

256 

8.1836 

.1209 

D 

Medium  long,  medium 
thick,  medium  rough 

264 

7.9924 

.1100 

E 

Long,  slim,  rough 

256 

8.7740 

.1182 

F 

Long,  thick,  rough 

270 

8.2741 

.1190 

G 

Short,  thick,  rough 

292 

8.2363 

.1111 

Composite 

1761 

8.1525 

.0452 

The  mean  yields  range  highest  in  Series  E,  F,  and  G,  and 
lowest  in  Series  A,  B,  C,  and  D.  But  between  the  highest  yield, 
Series  E,  and  the  lowest,  Series  B,  there  is  a difference  of  only 
1.06  ounces  of  shelled  grain  per  plant.  This  difference,  though 
small,  might  be  significant  did  not  the  yields  of  the  check  rows 
(Figure  1)  show  that  Series  E was  favored  by  a variation  in  the 
fertility  of  the  soil.  Doubtless  Series  F and  G were  likewise  fav- 
ored. 

There  were  then  no  significant  differences  between  the  yields 
of  plants  produced  from  the  various  seed-ears  representing  an 
extremely  wide  range  of  form  and  identation,  in  the  variety  Boone 
County  White. 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  46 


•Showing  the  comparative  location  of  Series  A to  G,  and  the  location  and  yield  in  pounds  of  grain  per  row 

of  the  check  rows  marked  O. 


Characters  Connected  with  the  Yield  of  the  Corn  Plant  9 


The  correlation  coefficients  determined  for  the  weight  of  shell- 
ed grain  as  the  subject  and  various  plant  characters  as  the  rela- 
tives are  now  shown. 


Table  II. — Correlations  Between  Variations  in  Plant  Characters  and 
Weight  or  Shelled  Grain  per  Plant. 


Character 

Coefficient  of 
correlation 

Probable 
error  (±) 

Days  from  planting  to  silking 

-.4181 

.0133 

Leaf  area  above  ear 

.0885 

.0167 

Leaf  area  below  ear 

.0565 

.0167 

Total  leaf  area 

.0702 

.0161 

Height  of  stalk 

.1109 

.0160 

Number  of  nodes  in  stalk 

.0843 

.0161 

Height  of  ear 

-.0006 

.0161 

Relative  position  of  the  ear-node 
Circumference  of  internode 

.0340 

.0161 

above  ground 

Circumference  of  internode 

.1846 

.0155 

above  ear 

.0893 

.0160 

Length  of  tassel 

-.1251 

.0170 

Number  of  tillers 

-.0160 

.0160 

Although  some  of  these  correlations  are  statistically  signifi- 
cant, none  of  them  are  high  enough  to  be  valuable  as  an  index  of 
yield.  The  negative  correlation  between  yield  and  the  age  of  the 
plant  at  silking,  the  highest  correlation  found,  would  doubtless 
vary  greatly  with  the  season. 

DISCUSSION 

The  results  of  this  brief  study  are  concordant  with  those  of 
other  studies  herein  cited,  in  finding  no  significant  correlations  be- 
tween the  yield  of  the  corn  plant  and  variations  in  its  visible  struc- 
tures and  characters.  But  these  and  all  similar  results  make  no 
proof  that  such  correlations  do  not  exist,  although  the  total  evi- 
dence has  come  from  a very  exhaustive  analysis.  To  accept  fully 
the  negation  of  correlations  would  lead  to  the  conclusion  that  the 
corn  plant  is  exhibiting  the  phenomenon  of  no  relationship  be- 
tween external  structure  and  function.  Of  course  the  correlations 
exist. 

Why  then  are  they  not  found  in  a measure  that  would  justify 


10  Missouri  Agr.  Exp.  Sta.  Research  Bulletin  46 

their  use  as  an  index  of  the  relative  ability  of  the  progeny  to 
yield?  A very  simple  explanation  may  be  suggested. 

In  all  studies  of  such  correlations  yield  has  been  treated,  con- 
sciously or  not,  as  a single  character  of  the  plant.  Obviously, 
this  conception  of  yield  is  fundamentally  wrong.  Yield  is  a per- 
formance, not  a character.  It  is  the  ultimate  performance  of  the 
whole  complex  relationship  of  functions  and  structures  that  make 
up  the  plant.  No  doubt  each  function  and  structure  varies  with 
the  environment.  No  doubt  each  variation  influences  yield ; but 
only  as  it  contributes  to  the  final  complex  result  of  all  variations. 
And  so  the  influence  of  a given  variation  upon  yield  cannot  be 
finally  measured,  simply  because  it  cannot  be  identified  and  sep- 
arated from  the  combined  influence  of  innumerable  other  variations. 

But  is  there  no  visible  index  of  yielding  ability  that  may  serve 
as  a guide  in  the  practical  operation  of  selecting  seed  corn?  It  was 
to  answer  this  question  that  all  correlation  studies  of  corn  were 
begun.  Certainly  there  is  such  an  index.  It  is  yield  itself — al- 
most the  old  and  simple  idea  of  selecting  the  biggest  ears. 

Taking  as  an  example  any  common  one-eared  variety  of  the 
Middle  West,  the  yield  of  grain  from  plant  to  plant  must  vary 
with  the  size  of  the  ear,  excluding  of  course  the  slight  variation 
in  shelling  percentage  and  the  losses  from  unsoundness.  So  far 
then  as  yield  can  be  improved  by  seed  selection,  the  most  exhaus- 
tive studies  have  discovered  no  better  method  than  field  selection 
of  the  biggest,  soundest  ears,  well  matured  and  unfavored  by  ap- 
parent differences  in  their  local  environment — stand,  fertility,  and 
so  forth.  Or  if  the  plant  bears  more  than  one  ear,  of  course  its 
total  yield,  rather  than  the  size  of  the  individual  ear,  should  be 
considered.  In  a given  environment  the  best  adapted  and  best 
yielding  strain  will  of  course  show  some  distinguishing  character- 
istic. For  example,  under  certain  conditions  the  highest  yielding 
strain  may  have  smooth  kernels.  But  it  does  not  follow  that  con- 
tinued close  selection  of  smooth  seed  ears  will  increase  or  even 
maintain  the  yield  of  the  strain.  For  by  that  operation  a specialized 
strain  not  so  well  balanced  with  the  environment  might  be  pro- 
duced. 


CONCLUSION 

Within  the  conventional  limits  of  a variety  of  corn,  no  varia- 
tion in  the  visible  structures  or  characters  of  a normal,  healthy 
plant  is  a reliable  index  of  the  relative  ability  of  its  progeny  to 


Characters  Connected  with  the  Yield  of  the  Corn  Plant  11 


yield.  The  relative  yield  of  the  mother  plant  is  the  only  indica- 
tion, uncertain  as  it  may  be,  of  the  relative  yield  of  the  progeny. 

This  conclusion  is  based  not  wholly  upon  the  brief  evidence 
presented  in  this  paper,  but  upon  the  total  evidence  contributed 
by  all  investigators  of  the  problem. 


BIBLIOGRAPHY 

1.  Duley,  F.  L.  and  Miller,  M.  F.  The  Effect  of  a Varying  Supply  of 
Nutrients  Upon  the  Character  and  Composition  of  the  Maize  Plant 
at  Different  Periods  of  Growth.  Mo.  Agr.  Expt.  Sta.  Res.  Bui.  42.  1921. 

2.  Montgomery,  E.  G.  Experiments  with  Corn.  Neb.  Agr.  Expt.  Sta. 

Bui.  112.  1909. 

3.  Hartley,  C.  P.  Producing  Higher  Yielding  Strains  of  Corn.  U.  S. 

Dept.  Agr.  Yearbook,  1909:  309-320. 

4.  Pearl,  R.  and  Surface,  F.  M.  Experiments  in  Breeding  Sweet  Corn. 

Me.  Agr.  Expt.  Sta.  Bui.  183.  1910. 

5.  Ewing,  E.  C.  Correlation  of  Characters  in  Corn.  Cornell  Univ.  Agr. 

Expt.  Sta.  Bui.  287.  1910. 

6.  Love,  H.  H.  The  Relation  of  Seed  Ear  Characters  To  Earliness  in 

Corn.  Amer.  Breeders  Ossoc.  Rpt.  8:  330-334.  1911. 

7 and  Wentz,  J.  B.  Correlations  Between  Ear  Char- 
acters and  Yield  in  Corn.  Jour.  Amer.  Soc.  Agron.,  8,  7:  315-322.  1917. 

8.  Sconce,  J.  H.  Scientific  Corn  Breeding.  Amer.  Breeders  Assoc.  Rpt. 

7:  43-50.  1911. 

9.  Funk,  E.  Ten  Years  of  Corn  Breeding.  Amer.  Breeders  Mag.  3,  4: 
295.  1911. 

10.  McCall,  A.  G.  and  Wheeler,  C.  S.  Ear  Characters  Not  Correlated 
with  Yield  in  Corn.  Jour.  Amer.  Soc.  Agron.,  5,  2:  117.  1913. 

11.  Williams,  C.  G.  and  Welton,  F.  A.  Corn  Experiments.  Ohio  Agr. 

Expt.  Sta.  Bui.  282.  1915. 

12.  Cunningham,  C.  C.  The  Relation  of  Ear  Characters  of  Corn  to 

Yield.  Jour.  Amer.  Soc.  Agron.,  8,  3:  188-196.  1916. 

13.  Hughes,  H.  D.  An  Interesting  Experiment  with  Seed  Corn.  Iowa 

Agr.,  17,  9:  424,  425,  428.  1917. 

14.  Hutcheson,  T.  B.  and  Wolfe,  T.  K.  Relation  Between  Yield  and  Ear 
Characters  in  Corn.  Jour.  Amer.  Soc.  Agron.,  10,  6:  250-225.  1918. 

15.  Olson,  P.  J.,  Bull,  C.  P.  and  Hayes,  H.  K.  Ear  Type  Selection  and 

Yield  in  Corn.  Minn.  Agr.  Expt.  Sta.  Bui.  174.  1918. 


12  Missouri  Agr.  Exp.  Sta.  Research  Bulletin  46 

II. — A Study  of  the  Relation  of  Certain  Ear 
Characters  to  Shelling  Percentage 
Shrinkage  and  Viability. 

This  study  was  made  in  1910  and  1911.  Its  purpose  was  to 
find  whether  variations  in  certain  characters  commonly  used  in 
judging  seed  ears  were  indicative  of  the  relative  shelling  per- 
centage, shrinkage  and  germination. 

MATERIAL  AND  GENERAL  METHODS 

In  1910,  660  sound  ears  of  a rough,  large-eared  strain  of 
Boone  County  White,  grown  on  rich  alluvial  soil,  harvested  in 
December  of  1909  and  stored  in  a tightly  boarded  crib  until  March 
1,  were  used  as  experimental  material.  They  will  be  designated 
as  Lot  A. 

In  1911,  500  sound  ears  of  the  same  variety,  but  of  a more 
variable  strain,  were  used.  They  too  had  been  grown  on  rich 
alluvial  soil,  but  had  been  harvested  early  in  October  and  air- 
dried  on  racks  in  a mouse-proof  seed  room  for  a period  of  12  weeks. 
They  will  be  designated  as  Lot  B. 

Both  lots  were  selected  at  random,  except  with  reference  to 
soundness.  In  both  lots  the  individual  ears  were  described  in  the 
details  hereafter  stated  in  Tables  I — IV.  All  descriptions  were 
recorded  by  the  same  person.  No  mathematical  correlations  were 
determined,  but  all  comparisons  were  made  between  two  classes 
showing  extreme  variation  of  the  character  in  question,  each  class 
constituting  about  15  percent  of  the  total  number  of  ears  in  the 
lot.  For  example,  in  studying  the  relation  of  length  of  ear  to 
shelling  percentage  in  Lot  A,  the  average  shelling  percentage  of 
the  100  longest  ears  was  compared  with  that  of  the  100  shortest 
ears  in  the  same  lot. 

THE  RELATION  OF  EAR  CHARACTERS  TO  THE  SHELL- 
ING PERCENTAGE  OF  THE  EAR 

In  Table  I is  shown  the  relation  of  various  ear  characters  to 
shelling  percentage,  as  determined  by  this  method.  The  differ- 
ences are  in  most  cases  slight  and  inconsistent.  Except  the  differ- 
ence between  light  and  heavy  ears,  which  may  be  attributed  to 
the  higher  moisture  content  of  the  latter  (see  Table  II),  the  size 


Characters  Connected  with  the  Yield  of  the  Corn  Plant  13 


and  shape  of  the  ear  show  no  significant  relation  to  shelling  per- 
centage; but  ears  marked  by  deep  kernels,  narrow  kernels,  and 
starchy  kernels  produced  a slightly  higher  proportion  of  grain 
than  ears  marked  by  shallow,  wide,  and  horny  kernels. 


Table  I. — Ear  Characters  and  Shelling  Percentage. 


Lot  A— 1910 

Lot  B- 

—1911 

Character  of 
the  ears 

Shelling 

percentage 

Ave.  weight 
of  grain 
(grams) 

Shelling 

percentage 

Ave.  weight 
of  grain 
(grams) 

Long 

84.1 

451.4 

84.4 

393.3 

Short 

86.1 

367.5 

84.5 

347.1 

Large  circumference 

84.4 

450.7 

84.7 

408.3 

Small  circumference 

84.8 

371.1 

86.3 

357.0 

Heavy 

83.8 

464.0 

86.0 

422.0 

Light 

89.9 

358.5 

85.7 

327.2 

Many  rows  of 

kernels 

85.0 

441.2 

85.5 

411.6 

Few  rows  of 

kernels 

83.3 

384.4 

84.5 

369.4 

Cylindrical 

84.1 

412.4 

85.2 

326.5 

Tapering 

84.4 

411.8 

84.6 

381.0 

Rough  indentation 

84.1 

421.7 

83.9 

396.8. 

Smooth  indentation 

84.2 

411.8 

84.2 

337.6. 

Deep  kernels 

85.7 

450.1 

86.4 

409.2 

Shallow  kernels 

83.1 

383.2 

83.2 

334.7 

Wide  kernels 

83.8 

415.8 

84.1 

382.2 

Narrow  kernels 

85.2 

419.0 

85.7 

393.0 

Horny  kernels 

82.0 

391.2 

83.8 

346.3 

Starchy  kernels 

85.5 

415.8 

85.6 

379.8 

THE  RELATION  OF  EAR  CHARACTERS  TO  THE  SHRINK- 
AGE OF  THE  EAR 

The  relation  of  ear  characters  to  shrinkage  was  studied  in 
Lot  B by  comparing  the  length,  circumference,  and  weight  of  the 
ears  as  first  stored,  with  their  length,  circumference  and  weight 
at  the  close  of  the  total  drying  period  of  6 weeks.  The  results  of 
this  study  are  shown  in  Table  II. 

Little  relation  is  shown  between  shrinkage  and  indentation 
or  between  shrinkage  and  the  length  and  shape  of  the  ear.  Ap- 
parently, however,  heavy  ears,  thick  ears,  deep-kerneled  ears,  and 
ears  with  a large  number  of  rows,  lost  considerably  more  weight 
than  ears  of  the  opposite  types. 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  46 


Table  II. — Ear  Characters  and  the  Average  Shrinkage  in  Length,  Cir- 
cumference and  Weight  of  Ears  of  Lot  B. 


Character 

Loss  in 

Length 

Loss  in  Circumference 

Loss  in 

Weight 

of  the  ears 

Inches 

Percent 

Inches 

Percent 

Grams 

Percent 

Long 

.4572 

4.4 

.3825 

5.3 

69.4 

15.6 

Short 

.3520 

4.0 

.3843 

5.2 

61.6 

16.5 

Large 

circumference 

.4250 

4.6 

.4624 

5.9 

75.4 

17.0 

Small 

circumference 

.3354 

3.4 

.3329 

4.9 

49.9 

12.6 

Many  rows 

.3790 

4.0 

.3930 

5.1 

77.3 

17.1 

Eew  rows 

.3612 

3.7 

.2295 

3.4 

59.8 

14.5 

Heavy 

.5140 

5.2 

.4183 

5.6 

92.4 

19.1 

Light 

.3412 

3.3 

.3300 

4.7 

44.5 

12.8 

Cylindrical 

.4362 

4.5 

.4000 

5.8 

63.9 

17.3 

Tapering 

.3710 

3.9 

.4000 

5.5 

65.8 

15.6 

Rough 

indentation 

.3651 

3.8 

.3475 

4.6 

58.2 

13.7 

Smooth 

indentation 

.4222 

4.5 

.4030 

5.9 

67.8 

14.2 

Deep  kernels 

.3849 

4.0 

.3520 

4.6 

89.7 

18.3 

Shallow  kernels 

.3108 

3.2 

.3700 

5.5 

59.8 

15.7 

In  the  same  lot  of  ears  the  rapidity  of  shrinkage  was  deter- 
mined by  weighing  at  intervals  of  two  weeks,  300  ears  grouped 
in  extreme  classes  as  previously  described. 

The  results  are  shown  in  Table  III. 


Table  III. — Ear  Characters  and  the  Progressive  Rate  of  Shrinkage. 


Character 


Percentage  of  Loss  in  weight 


of  the  ears 

2-wks 

4-wks 

6-wks 

8-wks 

10-wks 

12-wks 

Total 

Large  circumerence 

8.6 

5.6 

1.7 

1.1 

0.7 

0.5 

18.2 

Small  circumference 

6.6 

5.3 

1.6 

1.2 

0.4 

0.4 

15.5 

Heavy 

9.6 

6.2 

1.6 

1.1 

0.5 

0.5 

19.5 

Light 

6.9 

4.5 

1.8 

0.8 

0.3 

0.6 

14.9 

Many  rows 

8.6 

6.2 

1.5 

1.0 

0.6 

0.3 

18.2 

Few  rows 

7.8 

5.2 

1.9 

1.0 

0.8 

0.4 

17.1 

Rough  indentation 

7.8 

5.5 

1.7 

0.9 

0.4 

0.5 

16.8 

Smooth  indentation 

8.2 

5.9 

1.8 

0.9 

0.8 

0.6 

18.2 

Deep  kernels 

10.5 

5.9 

1.9 

0.9 

0.6 

0.2 

20.0 

Shallow  kernels 

7.9 

5.3 

1.5 

1.2 

0.5 

0.2 

16.6 

Horny  kernels 

8.4 

5.7 

1.6 

1.1 

0.2 

0.4 

17.4 

Starchy  kernels 

8.9 

5.7 

1.8 

1.0 

0.4 

0.5 

18.3 

Characters  Connected  with  the  Yield  of  the  Corn  Plant  15 


It  may  be  noted  first  that  in  all  classes  of  ears  more  than  75 
percent  of  the  total  shrinkage  occurred  during  the  first  four 
weeks,  and  that  thereafter  the  shrinkage  in  all  classes  of  ears  was 
very  slight  from  one  two-week  interval  to  another.  Weather  con- 
ditions were  about  the  average  for  October,  November  and  De- 
cember in  this  section.  These  results  then  may  indicate  the 
probable  time  required  to  air-dry  seed  corn  under  good  condi- 
tions of  farm  storage.  Apparently  it  would  not  be  necessary  to 
keep  the  seed  ears  on  racks  or  various  other  drying  devices  for 
longer  than  a month.  They  could  then  be  stored  in  a more  con- 
venient bulk  without  damage  because  of  the  moisture  they  con- 
tained. Their  remaining  moisture  would  be  given  off  very  slowly 
and  uniformly  over  a long  period. 

There  seems  little  significance  in  the  relative  rates  of  shink- 
age  by  ears  of  the  different  types.  Large  ears  and  heavy  ears 
lost  moisture  more  rapidly  during  the  first  two  weeks  than  ears 
of  the  opposite  types,  due  probably  to  their  large,  heavy  cobs. 
The  comparatively  rapid  drying  of  deep-kerneled  ears  may  indi- 
cate the  desirability  of  this  type  for  seed,  provided  they  are  also 
large,  sound,  and  well  matured. 


THE  RELATIONS  OF  EAR  CHARACTERS  TO  VIABILITY 

At  the  time  of  this  study  the  ears  (Lot  A)  were  two  years 
old.  They  had  been  harvested  in  December  1909  and  stored  for 
nearly  3 months  under  rather  poor  conditions  before  they  were 
sent  to  the  Experiment  Station.  Their  shelling  percentage  had 
been  determined  (Table  I)  and  the  grain  of  individual  ears,  stored 
separately  in  bottles,  had  been  fumigated  several  times  with  hydro- 
cyanic acid  gas.  In  November,  1911  this  seed  was  tested  for  germ- 
ination. 

To  make  the  tests,  kernels  were  planted  at  a depth  of  1 inch 
in  sand  which  was  kept  at  a temperature  of  about  80°F  during 
the  day  and  about  60°F  during  the  night,  and  in  a fairly  uniform 
condition  of  moisture.  A composite  hundred  kernels  from  each 
ear  of  the  GGO-ear  lot — a total  of  66,000  kernels — were  planted  in 
two  equal  series,  one  12  days  later  than  the  other.  Ten  days  after 
planting,  the  numbers  of  strong  sprouts,  weak  sprouts,  and  sprouts 
not  appearing  above  the  ground,  were  counted.  The  results  are 
given  in  the  following  table. 


16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  46 


Table  IV. — Ear  Characters  and  Germination. 
(Percentage  of  Germination  in  10  Days) 


Character 
of  the  ears 

Strong 

plants 

Weak  Plants  not 

plants  above  ground 

Total 

germination 

Long 

34.8 

12.1 

7.2 

54.1 

Short 

37.3 

13.3 

9.6 

60.1 

Large  circumference 

30.5 

11.0 

7.3 

48.8 

Small  circumference 

45.4 

12.9 

6.8 

65.1 

Heavy 

28.3 

11.8 

7.1 

47.2 

Light 

42.2 

12.6 

7.0 

61.8 

Many  rows  (22  and  more)  30.9 

11.5 

6.4 

48.8 

Few  rows  (16  and  less) 

44.7 

13.2 

7.0 

64.9 

Twisted  rows 

39.6 

12.6 

6.8 

59.0 

Straight  rows 

38.1 

12.0 

6.6 

56.7 

Cylindrical 

38.2 

11.8 

6.2 

56.2 

Tapering 

39.0 

12.4 

7.2 

58.6 

Close  spaced  rows 

42.4 

13.7 

7.5 

63.6 

Open  spaced  rows 

37.2 

11.6 

8.3 

57.1 

Rough  indentation 

35.3 

12.6 

7.5 

55.4 

Smooth  indentation 

46.4 

11.8 

5.4 

63.6 

Wide  kernels 

33.4 

12.5 

7.8 

53.7 

Narrow  kernels 

35.5 

12.1 

5.8 

53.4 

Deep  kernels 

27.0 

11.4 

6.7 

46.6 

Shallow  kernels 

44.2 

12.9 

5.5 

62.0 

Horny  kernels 

54.4 

8.0 

4.4 

66.8 

Medium  horny  kernels 

39.4 

11.8 

6.1 

57.3 

Starchy  kernels 

36.0 

10.1 

7.2 

53.3 

Large  germs 

34.1 

10.9 

6.4 

51.4 

Small  germs 

41.6 

12.4 

6.1 

60.1 

High  shelling  percentage 

30.2 

11.5 

6.5 

48.2 

Low  shelling  percentage  43.6 

11.3 

5.8 

60.7 

Heavy  grains 

32.3 

12.6 

7.4 

52.3 

Light  grains 

37.1 

11.9 

5.5 

54.5 

Heavy  cobs 

33.4 

12.2 

7.0 

52.5 

Light  cobs 

39.3 

12.5 

6.3 

58.1 

A brief  inspection  of  the  data  will  show  that  seed  from  short 
ears,  light  ears,  ears  with  few  rows,  and  ears  of  small  circumfer- 
ence, germinated  better  than  seed  from  ears  of  the  opposite  ex- 
treme types.  However,  it  can  hardly  be  assumed  that  these  var- 
ious characteristics  of  size  bear  a direct  relation  to  the  viability 
of  the  seed.  Each  of  them  is  in  some  degree  merely  an  expression 
of  the  circumference  or  weight  of  the  cob ; and  one  might  expect 
a comparatively  low  germination  in  seed  borne  on  a large,  sappy 


Characters  Connected  with  the  Yield  of  the  Corn  Plant  17 


cob,  because  of  the  unfavorable  effect  of  a higher  moisture  con- 
tent. Some  verification  of  this  is  found  in  the  fact  that  seed  from 
light  cobs  germinated  58  percent,  while  seed  from  heavy  cobs 
germinated  52  percent. 

The  data  do  not  show  a material  difference  in  the  germina- 
tion of  seed  from  ears  extremely  variable  in  shape  and  in  the  form 
and  spacing  of  the  kernel  rows.  However,  smooth,  shallow,  horny 
kernels,  germinated  better  than  rough,  deep,  and  starchy  kernels, 
respectively.  Small  germs  sprouted  better  than  large  germs. 

It  is  possible  that  the  treatment  of  the  seed  previous  to  the 
germination  tests — late  harvesting,  3 months  storage  in  a crib,  and 
several  fumigations  with  hydrocyanic  acid  gas — may  have  affect- 
ed differently  the  viability  of  the  various  types.  Certainly  the 
viability  of  all  types  was  very  low  as  a result  of  this  treatment. 

SUMMARY 

1.  Ears  extremely  characterized  by  deep  kernels,  narrow 
kernels  or  starchy  kernels,  had  a slightly  higher  shelling  percentage 
than  ears  of  the  opposite  extremes.  No  other  characteristics  of 
the  ear  showed  a significant  relation  to  the  proportion  of  grain. 

2.  Heavy  ears,  thick  ears,  deep-kerneled  ears,  and  ears  with 
a large  number  of  rows,  lost  considerably  more  weight  than  ears 
of  the  opposite  extremes,  during  a total  drying  period  of  6 weeks. 
These  characteristics  are  of  course  closely  related  to  the  size  of 
the  cob.  Other  characteristics  of  the  ear  showed  no  relation  to 
the  total  loss  of  moisture. 

3.  In  all  types  of  ears  more  than  75  percent  of  the  total 
shrinkage  occurred  during  the  first  4 weeks  of  a drying  period  of 
12  weeks.  Additional  shrinkage  was  very  slow  over  the  following 
8 weeks  period.  This  indicates  that  when  seed  corn  has  been  air- 
dried  on  racks  or  other  devices  for  about  a month,  under  climatic 
conditions  similar  to  those  of  this  experiment,  it  may  safely  be 
stored  in  a more  convenient  bulk. 

4.  Smooth  kernels,  shallow  kernels,  horny  kernels,  and  ker- 
nels with  small  germs,  showed  a higher  viability  than  kernels  of 
the  opposite  extremes.  No  characteristic  of  the  ear  as  a whole 
showed  a relation  to  viability  which  may  not  be  traced  to  the  mois- 
ure  content  of  the  cob.  Possibly  the  previous  treatment  of  the  seed 
influenced  the  relative  viability  of  the  different  types. 


UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 
AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  47 


Localization  of  the  Factors  Determining 
Fruit  Bud  Formation 


(Publication  Authorized  August  26,  1921) 


COLUMBIA,  MISSOURI 
SEPTEMBER,  1921 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 


EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  EANSING  RAY,  P.  E.  BURTON,  H.  J.  BEANTON, 

St.  Eouis  Joplin  Paris 


ADVISORY  COUNCIL, 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 

J.  C.  JONES,  Ph.  D.,  LL.  D„  ACTING  PRESIDENT  OF  THE  UNIVERSITY 


STATION  STAFF 
SEPTEMBER,  1921 


AGRICULTURAL  CHEMISTRY 

C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

VV.  S.  Ritchie,  A.  M. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  Sieveking,  B.  S.  in  Agr. 

AGRICULTURAL  ENGINEERING 

J.  C.  WoolEy,  B .S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

E.  F.  Hopkins,  Ph.  D. 

DAIRY  HUSBANDRY 

A.  C.  Ragsdale.  B.  S.  in  As*-- 
W.  W.  Swett,  A.  M. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride, 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  A.  M. 

O.  W.  Letson,  B.  S.  in  Agr. 

B.  M.  King,  B.  S.  in  Agr. 

A.  C.  Hill 

Miss  Bertha  C.  Hite,  A.  B.1 
Miss  Pearl  Drummond,  A.  A.1 

*In  service  of  U.  S.  Department  of  Agr 
2On  leave  of  absence. 


RURAL  LIFE 

O.  R.  Johnson,  A.  M. 

S.  D.  Gromer,  A.  M. 

E.  L.  Morgan,  A.  M. 

Ben  H.  Frame,  B.  S.  in  Agr. 


HORTICULTURE 

V.  R.  Gardner,  M.  S.  A. 

H.  D.  Hooker,  Jr.,  Ph.  D. 

J.  T.  Rosa,  Jr..  M.  S. 

F.  C.  Bradford,  M.  S. 

H.  G.  Swartwout,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY' 

H.  L.  Kempster,  B.  S. 

Earl  W.  Henderson 


SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M. 

W.  A.  Albrecht,  Ph.  D. 

F.  L.  DulEy,  A.  M.2 
R.  R.  Hudelson.  A.  M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr. 

Richard  Bradfield,  A.  B. 

O.  B.  Price,  B.  S.  in  Agr. 

VETERINARY  SCIENCE 

J.  W.  Connaway,  D.  V.  S.,  M.  D. 

L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 

OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Sercretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Jane  Frodsham,  Librarian 
E.  E.  Brown,  Business  Manager 

culture,  Seed  Testing  Laboratory. 


LOCALIZATION  OF  THE  FACTORS  DE- 
TERMINING FRUIT  BUD  FORMATION 

H.  D.  Hooker,  Jr.  and  F.  C.  Bradford 


Belief  in  a relationship  between  slow  growth  and  a fruitful 
condition  in  apple  and  pear  trees  has  come  down  to  the  present 
with  the  approval  of  many  generations  of  growers.  Said  John 
Lawrence1,  in  1717,  concerning  the  pear:  “*****but  yet  for  the 
sake  of  that  noble  Fruit  which  some  Kinds  produce  by  the  Help 
of  a Wall,  it  is  worth  while  to  humble  him  and  keep  him  in  Order. 
For  which  purpose****I  sometimes  plash  the  most  vigorous 
Branches,  cutting  them  near  the  place  from  whence  they  shoot, 
more  than  half  through,  which  effectually  checks  its  Vigour,  and 
consequently  renders  it  more  disposed  to  make  weaker  Shoots,  and 
form  bearing  Buds.” 

The  chief  concern  of  the  older  writers  on  fruit  bud  formation 
seems  to  have  been  the  prevention  of  excessive  growth.  This  was 
natural,  since  they  were  dealing  chiefly  with  fruit  gardens,  manured 
and  cultivated  and  consequently  with  trees  growing  luxuriantly. 
When  fruit  growing  spread  to  the  orchard  the  literary  heritage 
from  the  garden  survived  and  though  there  was  an  undoubted 
realization  of  the  unfruitfulness  of  greatly  weakened  trees  it  is 
but  recently  that  there  has  been  a crystallization  into  definite 
phrases  of  this  feeling  that  a certain  amount  of  growth  is  necessary 
for  fruit  bud  formation  and  that,  within  limits,  fruitfulness  and 
vegetative  development  are  associated  phenomena. 

SOME  OF  THE  FACTORS  INVOLVED 

The  work  of  Klebs2,  of  Fisher3,  of  Kraus  and  Kraybill4,  and 
of  Hooker5  has  given  some  conception  of  the  internal  chemical 
factors  connected  with  the  initiation  of  fruit  bud  differentiation. 
Briefly  stated,  this  seems  to  be  associated  primarily  with  carbohy- 
drate accumulation  and  in  apple  spurs,  with  starch  storage  in 
particular.  However,  even  though  carbohydrate  accumulation  oc- 
cur, fruit  and  differentiation  does  not  take  place  if  there  be  a lim- 
iting factor  which  seriously  retards  or  altogether  stops  vegetative 
growth.  The  inference  seems  warranted,  therefore,  that  the  supply 
of  water,  of  heat6,  of  nitrates  or  of  any  other  essential  nutrient 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  47 


may  so  check  growth  and  carbohydrate  utilization  that  carbohy- 
drate accumulation  results,  if  conditions  be  favorable  for  carbohy- 
drate manufacture ; any  one  of  these  factors  may  become  limiting 
and  prevent  fruitfulness,  though  under  field  conditions  the  nitrogen 
supply  seems  to  be  the  factor  most  frequently  operative  in  this 
direction.  When  the  nitrogen  supply  is  plentiful,  carbohydrate 
is  usually  found  in  the  plant  in  small  amounts,  because  it  has  been 
utilized  in  growth ; when  the  nitrogen  supply  is  low,  carbohydrate 
is  usually  found  accumulated  in  relatively  large  amounts.  In  the 
two  year  cycle  involving  fruit  bud  differentiation  one  year  and  fruit 
formation  the  next,  through  which  most  apple  spurs  on  fruitful 
trees  usually  pass,  starch  accumulation  is  associated  with  a rela- 
tively low  nitrogen  content  during  the  period  of  fruit  bud  differ- 
entiation the  one  year  and  a practical  absence  of  starch  is  associ- 
ated with  an  exceptionally  high  nitrogen  content  during  the  period 
of  fruit  setting  the  other  year.  ^ 

It  should  be  pointed  out  that  this  inverse  correlation  between 
nitrogen  and  carbohydrate  (particularly  starch)  content  does  not 
represent  a relationship  of  fundamental  importance  for  fruit  bud 
differentiation.  It  is,  in  a sense,  accidental,  though  it  is  common 
because  nitrogen  supply  is  most  often  the  limiting  factor  determin- 
ing carbohydrate  accumulation.  In  case  some  other  factor,  for  ex- 
ample water  supply,  were  operative  in  checking  growth,  it  is  clear 
that  carbohydrate  accumulation  might  take  place  even  in  the  pres- 
ence of  abundant  nitrogen.  In  fact  some  such  situation  must  ob- 
tain in  those  spurs  on  certain  apple  varieties  which  form  fruit 
buds  regularly  every  year.  If  fruit  setting  depend  on  the  presence 
of  a relatively  large  amount  of  nitrogen  in  the  spur  as  Harvey  and 
Murneek7  suggest  and  if  fruit  bud  differentiation  depend  on  starch 
accumulation,  then  large  amounts  of  nitrogen  and  of  carbohydrates 
must  be  present  almost  simultaneously  in  these  spurs.  This  situ- 
ation has  been  observed  in  spurs  of  Payne’s  Late  Keeper,  a local 
variety  in  which  a large  percentage  of  the  spurs  are  characterized 
by  successive  fruit  bud  formation.  A sample  of  bearing  spurs 
collected  July  3,  1920,  had  1.236  per  cent  nitrogen  and  3.16  per 
cent  starch.  Comparison  of  these  figures  with  the  data  published 
in  Research  Bulletin  40  of  this  Station  shows  that  this  nitrogen 
content  is  of  the  order  found  in  spurs  of  other  varieties  during  the 
spring  of  their  bearing  year  and  that  the  starch  content  is  equal 
in  amount  to  that  found  in  those  spurs  of  these  varieties  that  are 
differentiating  fruit  buds. 


Localization  of  Factors  Determining  Fruit  Bud  Formation  5 


THE  RELATION  OF  GROWTH  TO  PERFORMANCE 

The  fact  that  very  little  growth  and  very  vigorous  vegetative 
development  are  alike  unfavorable  for  fruit  bud  differentiation 
suggests  relationship  between  spur  growth  and  spur  performance 
in  the  apple.  Roberts8  found  that  in  Wealthy  and  other  apple 
varieties  under  certain  conditions  spurs  of  certain  length  growths 
showed  the  highest  percentage  of  fruit  bud  differentiation  and  that 
both  longer  and  shorter  spurs  showed  lower  percentages. 

To  establish,  for  closer  selection  of  samples  for  chemical  study, 
the  value  of  such  an  index  to  the  probable  performance  of  the  in- 
dividual spurs  under  conditions  obtaining  in  the  trees  growing  in 
<the  University  orchard  at  Columbia  studies  of  spurs  of  several 
varieties  were  undertaken.  Measurements  totaling  around  13,000 
were  made,  work  proceeding  with  each  variety  till  several  succes- 
sive series  showed  no  change  in  the  results  obtained.  For  purposes 
of  this  investigation  no  spur  was  considered  which  had  not  blos- 
somed at  least  once;  growth  in  one  year  of  over  10  cm.  was  arbi- 
trarily considered  to  remove  the  twig  from  the  spur  class  to  the 
shoot  class.  The  massed  results  are  shown  in  Table  1,  in  terms 
of  percentage  of  spurs  in  each  class  forming  fruit  buds.  The  figures 
presented  here  are  from  purely  vegetative  growths,  i.  e.,  no  meas- 
urements of  growth  of  any  spur  in  its  blossoming  season  are  in- 


Table  1. — Percentage  of  Spurs  of  Various  Length  Classes  Which  Formed 

Fruit  Buds 


Length 

(cm.) 

Gano 

Jonathan 

Devonshire 

Duke 

Wealthy 

0.1 -0.5 

17.5 

39.3 

15.3 

49.7 

0.6- 1.0 

45.7 

53.8 

17.5 

62.9 

1. 1-1.5 

60.6 

57.0 

11.9 

62.5 

1. 6-2.0 

69.2 

76.8 

30.0 

76.3 

2.1-3.0 

65.3 

60.4 

25.0 

62.8 

3. 1-4.0 

74.6 

72.7 



64.8 

4.1-5.0 

70.7 

63.3 



70.0 

5. 1-6.0 

79.2 

78.1 

50.0 

61.1 

6. 1-7.0 

88.6 

70.0 



81.2 

7. 1-8.0 

88.7 

100.0 

66.7 

8. 1-9.0 

80.5 

81.3 

66.7 

9.1-10.0 

74.0 

68.4 

— 

75.0 

3.0-10.0 

77.3 

75.2 

15.5 

69.5 

Average 

54.4 

53.4 

16.4 

60.2 

6 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  47 


eluded.  It  may  be  stated,  however,  that  inclusion  of  the  growths 
during  a season  of  fruiting  made  no  material  difference  except  in 
the  percentage  of  fruit  buds  formed ; the  relative  positions  of  the 
various  classes  remained  the  same. 

These  figures  show,  in  each  variety,  an  increase  in  the  per- 
centage of  fruit  buds  formed,  with  an  increase  in  the  growth  of  the 
current  year.  In  some,  however,  the  rise  from  the  lowest  class  to 
the  next  higher  is  much  more  abrupt  than  in  others,  in  Gano  from 
17.5  to  45.7  as  compared  with  a rise  from  39.3  to  53.8  per  cent  in 
Jonathan.  Viewing  it  in  another  way:  in  the  lowest  growth  class 
Gano  formed  less  than  half  the  percentage  of  fruit  buds  that  Jona- 
than spurs  making  the  same  growth  formed.  This  difference  could 
not  be  due  to  the  Jonathan  growths  averaging  nearer  the  upper 
limit  of  the  class  than  did  the  Gano,  for  actually  conditions  were 
reversed.  It  seems  quite  evident  that  Jonathan  will  form  a great- 
er percentage  of  fruit  buds  on  very  short  growths  than  will  Gano. 

Were  spurs  defined  as  growths  up  to  3 cm.  only,  there  would 
be,  in  all  four  varieties,  a maximum  percentage  of  fruit  bud  forma- 
tion in  those  growths  between  1.6  and  2.0  cm.  However,  in  each 
case,  except  Devonshire  Duke,  the  percentage  of  fruit  buds  formed 
on  growths  between  3.1  and  10.0  cm.  is  higher  than  in  any  of  the 
smaller  classes.  In  other  words,  there  is  a strong  tendency,  in  the 
trees  examined,  for  a continuing  increase  in  percentage  of  fruit 
bud  formation  with  increase  in  growth. 

Table  2 shows  the  percentage  distribution  of  these  same  spurs 


Table  2.— 

-Percentage  oe  Total 

Number  oe 
Class. 

Spur  Growths  in 

Each  Length 

Length 

(cm.) 

Gano 

Jonathan 

Devonshire 

Duke 

Wealthy 

0. 1-0.5 

6.7 

24.2 

40.6 

25.1 

0.6-1. 0 

17.4 

33.2 

45.5 

32.3 

1. 1-1.5 

14.0 

13.4 

7.0 

13.8 

1. 6-2.0 

9.3 

7.6 

5.6 

7.4 

2.1-3.0 

11.7 

5.9 

0.7 

9.5 

3. 1-4.0 

9.7 

4.0 

0.0 

3.3 

4. 1-5.0 

8.2 

1.9 

0.0 

2.3 

5. 1-6.0 

5.7 

3.2 

0.7 

1.6 

6. 1-7.0 

6.7 

2.1 

0.0 

1.8 

7. 1-8.0 

4.5 

1.7 

0.0 

1.6 

8. 1-9.0 

2.7 

1.3 

0.0 

0.8 

9.1-10 

3.5 

1.3 

0.0 

1.1 

Localization  of  Factors  Determining  Fruit  Bud  Formation  7 


in  the  various  length  classes.  In  each  variety  there  is  a larger  per- 
centage of  spurs  in  the  0. 6-1.0  cm.  class  than  in  any  other.  The 
close  parallel  in  the  distribution  of  Jonathan  and  of  Wealthy  spurs 
is  striking.  Gano  and  Devonshire  Duke  have  different  curves  of 
distribution. 


ANALYSIS  OF  DATA 

Correlation  between  spur  growth  and  performance  implies  a 
considerable  degree  of  autonomy  in  the  spur.  If  this  quasi  inde- 
pendence be  great,  then  studies  of  the  factors  affecting  fruit  bud 
formation  may  well  be  focused  on  the  spur,  taking  little  heed  of 
the  remainder  of  the  tree.  Because  of  the  bearing  of  this  matter 
on  other  investigations  under  way  the  data  accumulated  from 
measurements  were  subjected  to  closer  analysis. 

By  Years. — Table  3 shows  the  percentages  of  fruit  bud  forma- 
tion in  different  years  in  spurs  selected  at  random  from  six  Gano 
trees  in  the  University  orchard  at  Columbia,  bearing  regularly  in 
the  odd  years.  Since  these  measurements  were  taken  in  the  win- 
ter of  1919-1920  no  figures  from  subsequent  years  for  these  spurs 
are  available.  Performances  in  the  bearing  year  of  any  spur  were 
not  considered ; consequently  all  these  figures  apply  to  spurs  hav- 
ing full  opportunity,  so  far  as  they  were  concerned  individually,  to 
form  fruit  buds.  This  they  did  abundantly  in  some  years — the  off 
years — and  very  meagerly  in  others — the  bearing  years. 

The  marked  difference  between  the  distribution  of  spur  per- 


Table  3. — Percentages 

or  Fruit  Bud  Formation 
Lengths,  in  Gano. 

in  Spurs 

or  Dieeerent 

Spur  length 

1915 

1916 

1917 

1918 

(cm.) 

0.1 -0.5 

57 

3 

38 

1 

0.6-1.0 

81 

4 

53 

10 

1. 1-1.5 

87 

17 

59 

28 

1. 6-2.0 

94 

0 

72 

0 

2.1-3.0 

93 

20 

58 

37 

3. 1-4.0 

95 

0 

71 

100 

4.1-5.0 

94 

50 

73 

0 

5. 1-6.0 

100 

50 

70 

0 

6.1-7.0 

95 



80 

7. 1-8.0 

100 

__ 

81 

8. 1-9.0 

100 

0 

83 

0 

9.1-10.0 

88 

0 

87 

— 

8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  47 


formance  in  the  bearing  years  of  Gano  (1915  and  1917)  and  in  the 
off  years  (1916  and  1918),  as  shown  in  Table  3,  indicates  that  the 
condition  of  the  whole  tree  influences  spur  performance  in  no  small 
measure.  The  figures  recorded  in  Table  1 for  Devonshire  Duke 
show  much  the  same  type  of  distribution  of  fruit  bud  differentia- 
tion as  is  shown  by  the  off  years  for  Gano  and  the  low  average 
percentage  of  fruit  bud  formation  in  the  Devonshire  Duke  suggests 
an  association  between  the  condition  of  the  tree  and  the  character 
of  the  distribution  curve. 

By  Trees. — Interesting  comparisons  between  spur  perform- 
ances on  two  Wealthy  trees,  standing  side  by  side,  are  shown  in 
Table  4.  One  bears  biennially  in  a pronounced  manner;  the  other 


Table  4. — Percentage  oe  Fruit  Buds  Formed  by  Spurs  oe  Various  Growths 
on  a Biennially  and  on  an  Annually  Bearing  Wealthy  Tree. 


Biennial 

Annual 

Growth 

1916 

1917 

1918 

1919 

1920 

1916 

1917 

1918 

1919 

1920 

(cm.) 

0. 1-0.5 

88.4 

6.7 

86.7 

0 

100.0 

53.5 

26.3 

62.5 

78.0 

73.2 

0.6-1. 0 

75.6 

0 

95.0 

0 

99.2 

25.8 

38.4 

74.0 

82.9 

86.9 

1.1-1. 5 

83.3 

0 

85.0 

0 

95.7 

20.0 

44.4 

69.4 

65.0 

92.6 

1. 6-2.0 

80.0 

0 

92.9 

0 

100.0 

28.6 

66.7 

63.6 

80.0 

100.0 

2. 1-3.0 

88.5 

0 

73.7 



96.8 

33.3 

28.6 

58.3 

66.7 

100.0 

3. 1-4.0 

85.5 

0 

70.0 

0 

100.0 

50.0 

60.0 

63.6 

100.0 

100.0 

4.1-5.0 

86.0 

0 

83.0 



100.0 

50.0 

60.0 

66.7 

100.0 

100.0 

5. 1-6.0 

66.7 

0 

100.0 



100.0 



57.2 

50.0 

80.0 

100.0 

6. 1-7.0 

75.0 

0 

100.0 



100.0 

33.3 

85.7 

100.0 

100.0 

100.0 

7. 1-8.0 

100.0 



100.0 



100.0 

50.0 

28.6 

33.3 

100.0 

100.0 

8. 1-9.0 





100.0 







80.0 

33.3 

100.0 



9.1-10.0 

100.0 

— 

100.0 

— 

50.0 

62.5 

— 

100.0 

100.0 

has  borne  with  considerable  regularity  each  year.  Here  again  there 
appears  a tendency  toward  mass  behavior,  to  a considerable  degree 
independent  of  the  growth  of  the  individual  spur.  In  the  biennial 
bearing  tree  this  tendency  is  naturally  the  more  pronounced ; in 
1917  almost  no  fruit  buds  were  formed  by  any  of  the  spurs  studied, 
regardless  of  the  growth  they  made,  and  in  1919  there  were  prac- 
tically no  spurs  to  be  considered,  for  nearly  all  spurs  studied  were 
bearing.  The  apparent  discrepancy  between  the  1918  and  1919 
figures  is  explained  by  a few  instances  of  consecutive  bearing. 
Again  it  is  emphasized  that  no  measurements  taken  for  the  bear- 


Localization  of  Factors  Determining  Fruit  Bud  Formation  9 


ing  year  of  any  spur  are  included ; these  figures  apply  in  every 
instance  to  non-bearing  spurs. 

By  Branches. — The  data  in  Table  5 are  fairly  representative 
of  conditions  on  the  annually  bearing  Wealthy  and  show  clearly 
that  its  annual  bearing  is  due  in  large  measure  to  individuality  in 
the  behavior  of  the  branches.  Not  all  branches  show  the  same 
uniformity  exhibited  by  those  selected ; nevertheless  there  is  in 
every  case  a pronounced  tendency  to  bear  or  not  to  bear. 


Table  5. — Performance  Records  oe  Individual  Limbs  on  an  Annually  Bear- 
ing Wealthy  Tree. 


Branch 

No.  spurs 
examined 

No.  spurs  blossoming 

1917 

1918 

1919 

1920 

A 

21 

0 

18 

0 

21 

B 

14 

12 

0 

14 

0 

C 

43 

2 

33 

1 

43 

D 

30 

2 

0 

30 

0 

Table  6 records  performances  of  individual  spurs  on  two 
branches  of  the  same  tree,  one  branch  alternating  with  the  other. 
There  is  little  relation  between  growth  and  fruit  bud  formation  in 
such  cases  as  these,  though,  to  be  sure,  the  very  short  growths 
are  few.  Spurs  240,  241,  243  and  123,  which  failed  to  form  fruit 
buds  with  the  others,  did  not  form  them  for  the  off  year.  This 
occurrence,  though  not  invariable,  appears  to  be  more  common 
than  the  opposite  and  points  to  a lack  of  complete  independence. 

Two  branches,  each  of  half-inch  diameter,  arising  at  points 
six  inches  apart  on  the  same  limb  of  a Jonathan  tree,  yielded  the 
data  recorded  in  Table  7.  Every  spur  is  included.  These  were 
young  branches  selected  at  random ; several  of  the  spurs  on  Branch 
A arose  on  1917  wood.  Neither  of  these  limbs  has  borne  biennially 
and  in  both  cases  the  spur  growth  of  the  year  preceding  bearing 
is  greater  than  that  of  the  year  which  was  not  characterized  by 
fruit  bud  formation.  Nevertheless,  large  growth  in  the  first  of  the 
two  vegetative  years,  as  exemplified  in  spurs  A8(1918)  and  B7- 
(1919)  did  not  result  in  fruit  bud  formation,  though  in  the  follow- 
ing year  very  short  growths,  as  represented  by  A2(1919)  and  B6- 
(1920)  were  accompanied  by  fruit  bud  formation. 

By  Year  of  Origin. — Uncomplicated  by  previous  history,  spurs 
in  their  first  year  as  individuals,  if  they  be  autonomous,  should 


10 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  47 


present  in  their  performance  some  indication  of  that  condition. 
If,  on  the  other  hand,  they  depend  somewhat  on  conditions  farther 
back  in  the  tree,  more  fruit  bud  formation  should  occur  in  spurs 
arising  during  the  off  year.  Table  8 shows  the  first  year  per- 
formance of  all  Wealthy  spurs  measured.  Under  conditions  pre- 
sented here,  spurs  arising  during  the  off  year  are  likely  to  form 
fruit  buds  regardless  of  their  growth  and  those  arising  during  the 
bearing  year  are  unlikely  to  form  fruit  buds,  regardless  of  their 
growth. 

Another  way  of  stating  essentially  the  same  fact  is  shown  by 
the  arrangement  used  in  Table  9.  This  shows  the  date  of  the 
initial  blossoming  of  all  spurs  studied.  Were  the  spur  completely 
autonomous  there  should  be  a uniform  yearly  rate;  the  marked 
alternation  actually  shown  is  therefore  particularly  significant. 

By  Departures  from  Alternation. — In  the  cases  of  successive 
blossoming  observed  in  Wealthy,  the  second  blossoming  occurred 
in  the  bearing  year  in  71  per  cent  of  the  total  number.  This  means 


Table  6. — Performance  Records  of  Individual  Spurs  on  Two  Limbs  of  an 
Annually  Bearing  Wealthy  Tree. 


( Growth  in  cm.  F — blossoming,  L = non-blossoming ) 


Spur  No. 

1918 

1919 

1920 

1921 

Spur  No. 

1918 

1919 

1920 

1921 

230 

.7 

F 

5.3 

F 

109 

F 

.7 

F 

L 

231 

1.2 

F 

2.0 

F 

110 

F 

.7 

F 

L 

232 

6.7 

F 

3.2 

F 

111 

F 

.3 

F 

L 

233 

.8 

F 

.8 

F 

112 

F 

.8 

F 

L 

234 

1.5 

F 

.7 

F 

113 

F 

.6 

F 

L 

235 

1.5 

F 

2.9 

F 

114 

F 

.4 

F 

L 

236 

1.2 

F 

.9 

F 

115 

F 

.4 

F 

L 

237 

1.5 

F 

.6 

F 

116 

F 

.8 

F 

L 

238 

1.3 

F 

1.7 

F 

117 

F 

.8 

F 

L 

239 

.5 

F 

3.3 

F 

118 

F 

.5 

F 

L 

240 

1.0 

1.1 

2.6 

F 

119 

F 

1.5 

F 

L 

241 

.8 

.8 

1.8 

F 

120 

F 

9.8 

F 

L 

242 

1.1 

F 

1.1 

F 

121 

F 

5.3 

F 

L 

243 

.6 

1.7 

6.2 

F 

122 

F 

1.7 

F 

L 

244 

.5 

F 

.2 

F 

123 

8.6 

1.5 

F 

L 

245 

4.0 

F 

3.4 

F 

124 

F 

2.4 

F 

L 

246 

5.3 

F 

.8 

F 

125 

F 

1.6 

F 

L 

247 

.5 

F 

2.5 

F 

126 

F 

9.0 

F 

L 

248 

.5 

F 

.9 

F 

127 

F 

1.7 

F 

L 

249 

2.3 

F 

1.1 

F 

128 

F 

1.1 

F 

L 

250 

.3 

F 

.5 

F 

129 

F 

6.6 

F 

L 

Localization  of  Factors  Determining  Fruit  Bud  Formation  11 


that  the  spurs  blossoming  in  the  off  year  have  much  better  chances 
of  forming  fruit  buds  immediately  than  those  that  blossom  in  the 
bearing  year. 

Still  further  evidence  of  a general  influence  affecting  fruit 
bud  formation  lies  in  the  growths  of  non-bearing  spurs  in  a crop 
year  as  compared  with  those  made  in  an  off  year.  If  the  general 
draft  of  the  crop  have  any  effect  it  should  be  reflected  in  the  growth 
of  the  non-bearing  spurs  at  that  time.  Data  from  Gano,  Jonathan 
and  Wealthy  show  an  almost  invariable  increase  and  decrease  of 
vegetative  growth  inversely  with  the  crop  (see  Table  10).  It  is 
least  pronounced  in  Jonathan,  where  the  alternation  of  crops  is 
less  pronounced.  Comparison  between  varieties  shows,  as  for 
example  between  1918  and  1919,  that  the  growths  of  two  varieties 
go  upward  and  that  of  the  other  downward,  in  keeping  with  the 
crops,  precluding  the  possibility  of  weather  influences  controlling 
this  increase  and  decrease. 

Unsupported,  these  data  would  be  open  to  the  objection  that 
the  spurs  not  bearing  in  the  crop  year  are  chiefly  barren  spurs 
whose  growth,  always  small,  is  submerged  by  the  figures  for  the 
prolific  spurs  in  the  off  year  but  constitutes  nearly  all  the  figures 

Table  7. — Performance  Records  of  Individual  Spurs  on  Two  Branches 
Arising  From  the  Same  Limb  on  a Jonathan  Tree. 

( Growth  in  cm.  F = blossoming,  L = non-blossoming ) 


1918 


1919 


1920 


1921 


Branch  A 1 


.5 


4.3 
.2 

1.4 
1.0 
2.6 

.7 

1.6 

3.7 

.5 

.7 

.5 

.8 

.3 

.3 

2.0 

.5 

.4 

.4 


F 

F 

F 

F 

F 

F 

F 

F 

1.0 

.6 

1.7 

1.7 

.5 

.3 

6.3 

4.0 
.4 

1.0 


L 

L 

L 

F 

L 

L 

L 

L 

F 

F 

F 

F 

F 

F 

F 

F 

F 

F 


2 

3 

4 

5 

6 

7 

8 


.3 

.3 

.5 

.2 

.6 

1.0 


Branch  B 1 


2 

3 

4 

5 

6 

7 

8 
9 
10 


F 

F 

F 

F 

F 

F 

F 

F 

F 


12 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  4 7 


for  the  crop  year.  Though  no  spur  which  had  not  blossomed  at 
least  once  was  recorded,  as  an  additional  check,  all  cases  in  which 
a spur  after  once  blossoming  had  failed  to  follow  the  usual  alter- 
nation were  tabulated  for  Gano. 

If  there  is  any  general  influence  affecting  fruit  bud  formation 
it  should  be  more  effective  toward  bringing  these  barren  spurs 
back  to  bearing  in  the  crop  year  than  in  the  off  year.  If,  in  addi- 
tion, consecutive  barrenness  arise  from  failure  to  bear  in  the  crop 
year,  this  same  general  influence  should  act  to  prevent  fruit  bud 
formation  for  blossoming  in  the  off  year.  Consequently  there 
should  be  more  cases  of  three-  and  five-year  successions  of  sterility 
than  of  two-  and  four-year  respectively.  If,  on  the  other  hand, 
there  is  no  general  influence,  the  frequency  distribution  should  be 


Table  8. — Percentage  oe  Fruit  Bud  Formation  on  Wealthy  Spurs  During 

Their  First  Year. 


Growth  (cm.) 

0.1- 

0.5 

0.6- 

1.0 

1.1- 

1.5 

1.6- 

2.0 

2.1-  3.1-  4.1- 
3.0  4.0  5.0 

5.1- 

6.0 

6.1- 

7.0 

7.1- 

8.0 

8.1- 

9.0 

9.1- 

10.0 

Biennial  Bearing  Tree 

Off  Year 

70 

71 

69 

80 

92  100  50 

100 

75 

100 

67 

Bearing  Year 

5 

0 

0 



0 0 __ 



0 





0 

Regular  Bearing  Tree 

Off  Year 

(for  branch) 

83 

91 

100 

0 

100  50  100 

100 

100 

Bearing  Year 
(for  branch) 

8 

0 

0 

0 

0 

0 

0 

Combined 

Off  Year 

76 

76 

76 

57 

93  75  67 

100 

83 

100 

75 

Bearing  year 

6 

0 

0 

0 

0 0 __ 

0 

0 

0 

0 

0 

in  uniformly  descending  order,  the  two-year  cases  being  most  fre- 
quent. The  actual  distribution  found  was:  two  years,  37;  three 
years,  115 ; four  years,  6 ; five  years,  9. 

Still  closer  analysis  lends  further  support.  The  two-  and  four- 
year  cases  originated  practically  equally  from  bearing  in  the  off  year 
and  from  failure  to  bear  in  the  crop  year.  Consequently,  when  com- 
bined and  averaged  they  have  little  significance.  However,  with  the 
growth  segregated  into  crop  year  and  off  year,  that  in  the  crop  year 
is  found  to  average  1.97  cm.  and  in  the  off  year  2.30  cm.  The  three- 
and  five-year  cases  originated  almost  entirely  from  failure  to  bear 
in  the  crop  year.  Therefore  their  growths,  if  there  be  assumed  a 
general  influence  outside  the  individual  spurs,  should  show  alter- 


Localization  of  Factors  Determining  Fruit  Bud  Formation  13 


nate  decreases  and  increases,  inverse  to  the  crop  of  the  tree.  In 
the  three-year  cases,  starting  with  the  off  year,  the  average  growths 
were  respectively  2.64;  0.91  and  1.37  cm.;  for  the  five-year  cases 
the  average  growths  were,  respectively:  1.1,  1.0,  1.1,  0.5,  and  0.7, 
showing  in  both  classes  a rhythm  inverse  to  the  crop.  In  all  cases, 
the  growth  averages  less  in  the  crop  year. 

A still  more  rigid  test  is  secured  by  assembling  cases  of  con- 
secutive unfruitfulness  and  averaging  their  growths  with  all 
growths  accompanying  fruit  bud  formation  eliminated ; these  fig- 
ures represent  purely  vegetative  growths  resulting  only  in  leaf  bud 
formation.  For  the  consecutive  years  of  most  marked  crop  alter- 


TablE  9. — Percentages  of  Initial  Blossoming  by  Years  in  Spurs  of  Gano, 
Wealthy,  and  Jonathan. 


Gano 

Wealthy 

J onathan 

1906 

1.2 

0.2 

1907 

0.6 

_ 

0.9 

1908 

0.8 



1.6 

1909 

1.0 



1.6 

1910 

3.8 



2.9 

1911 

1.8 

0.8 

6.8 

1912 

14.8 

0.8 

9.5 

1913 

3.6 

3.0 

16.5 

1914 

22.6 

0.4 

9.5 

1915 

3.2 

3.4 

19.6 

1916 

40.2 

1.1 

7.1 

1917 

0.4 

49.1 

17.1 

1918 

5.8 

0.4 

6.0 

1919 

0.2 

35.4 

1.3 

1920 



0.0 



1921 

— 

5.8 

— 

nation  these  averages,  beginning  with  the  off  year,  are  respective- 
ly: 1.19  cm.,  0.96  cm.,  1.15  cm.  and  0.89  cm.  If  the  last  year  of  the 
unfruitful  succession  be  included — the  year  before  resumption  of 
bearing — these'  differences  are  accentuated,  viz.:  1.66  cm.,  1.03  cm., 
3.28  cm.  and  0.88  cm.  However,  even  with  these  omitted,  the  ef- 
fect on  the  purely  vegetative  growth  is  interesting. 

An  Annually  Bearing  Limb. — To  complete  the  statistical  an- 
alysis, data  from  certain  spurs  on  one  limb  of  the  annually  bearing 
Wealthy  are  assembled  in  Table  11.  These  spurs  have  histories 
tracing  back  at  least  to  1914;  most  of  them  are  older.  This  limb 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  4 7 


is  characterized  by  a tendency  to  bear  annually.  That  this  is  an 
acquired  tendency  is  shown  by  the  increase  from  1914  to  1919  in 
the  number  of  spurs  blossoming,  a rise  from  one  in  the  former  year 
to  nine  in  the  latter.  The  proportion  of  cases  of  fruit  bud  forma- 
tion in  successive  years  is  unusual  for  this  variety.  Even  here 
the  relationship  between  growth  and  fruit  bud  formation  is  not 
sharply  defined  and  is  revealed  only  by  a general  tendency  toward 
increased  growth  as  the  spurs  approached  a fruitful  condition. 


Table  10.— Growth  oe  Non-Blossoming  Spurs  in  Relation  to  the  Percentage 
oe  Spurs  Blossoming  in  the  Same  Year. 


1920 

1919 

1918 

1917 

1916 

1915 

1914 

Gano 

Growth  (cm.) 



1.91 

0.85 

3.23 

1.27 

2.79 

1.15 

Crop  (per  cent) 



3-8 

64.4 

1.2 

79-3 

6.2 

55.3 

Jonathan 

Growth  (cm.) 



0.81 

0.76 

1.54 

1.96 

1.20 

1.35 

Crop  (per  cent) 



14.4 

3i.5 

55-2 

21.3 

55-0 

24.2 

Wealthy 

Growth  (cm.) 

1.26 

1.21 

1.45 

1.19 

1.79 

0.74 

1.23 

Crop  (per  cent) 

0-3 

89.7 

0.3 

65.2 

2.1 

9 4 

2.6 

DISCUSSION 

Apparently,  then,  the  data  presented  show,  on  the  one  hand 
a relation  between  spur  growth  and  performance  and  on  the  other 
hand  little  or  none  of  such  relation.  However,  they  are  not  irre- 
concilable. They  reflect  different  conditions.  Under  certain  con- 
ditions practically  all  spurs  either  form  fruit  buds  or  they  do  not; 
under  other  conditions  some  spurs  form  fruit  buds  while  others, 
intermixed  with  them,  do  not.  In  the  latter  case  the  correlation 
between  growth  and  fruit  bud  formation  attains  some  importance. 
Spurs  showing  several  successive  years  of  unfruitfulness  have  a 
tendency  toward  increased  growth  in  the  year  of  fruit  bud  forma- 
tion. This  accounts  for  a little  more  of  the  correlation.  Combined 
in  massed  figures  these  two  tendencies  establish  the  correlation  as 
shown  in  Table  1. 

However,  the  spurs  that  are  making  successive  vegetative 
growths  show  a tendency  toward  uniformity  even  in  the  amount 
of  this  growth,  as  shown  in  Table  7.  This  suggests  strongly  that 
the  relationship  between  growth  and  performance  is  a parallel 


Localization  of  Factors  Determining  Fruit  Bud  Formation  15 


manifestation  of  the  same  or  related  influences  and  not  in  itself 
a cause  and  effect  relationship. 

The  succession  of  units  possible  in  performance  is  most  strik- 
ing. On  the  one  extreme  is  the  tree  (Table  4)  ; next  smaller,  the 
scaffold  limb  (Table  6)  ; still  in  descending  order,  the  branch 
(Table  7)  and  finally- the  spur  (Table  11).  This  points  to  the 
probable  importance  of  influences  farther  back  in  the  tree  than  the 
spurs  in  promoting  fruit  bud  formation.  The  influence  must  be 
strong  in  some  cases,  causing  general  uniformity,  either  in  fruit- 
fulness or  in  unfruitfulness.  In  other  cases  it  may  be  less  positive 

Table  11. — Performance  Record  oe  Individual  Spurs  on  a Branch  oe  an 
Annually  Bearing  Wealthy  Tree. 


( Growth  in  cm.  F — blossoming,  L — non-blossoming ) 


Spur  No. 

1914 

1915 

1916 

1917 

1918 

1919 

1920 

1921 

169 

.7 

1.3 

6.2 

.8 

F 

.6 

F 

L 

170 

1.2 

2.3 

9.1 

F 

3.8 

F 

1.0 

F 

172 

.6 

F 

9.4 

8.0 

3.5 

1.3 

1.4 

F 

173 

1.0 

.9 

F 

7.9 

4.2 

F 

.2 

L 

174 

.7 

1.4 

F 

2.1 

.8 

F 

.5 

F 

177 

1.2 

1.0 

6.1 

F 

F 

F 

.7 

F 

178 

.5 

4.9 

.5 

3.9 

.6 

F 

1.1 

F 

180 

.5 

.6 

6.2 

6.4 

F 

1.0 

1.3 

F 

181 

.5 

.5 

.7 

.6 

.5 

F 

.8 

F 

183 

.2 

.2 

.4 

.3 

F 

F 

F 

L 

184 

.5 

1.1 

2.0 

5.7 

1.5 

F 

.4 

L 

186 

.4 

.4 

.7 

.8 

F 

F 

.3 

L 

187 

F 

.5 

3.1 

9.3 

F 

.4 

F 

L 

188 

.9 

.4 

.5 

F 

.4 

.5 

F 

L 

191 

.5 

.6 

2.5 

1.1 

F 

.8 

F 

F 

and  the  performance  of  the  individual  spur  determined  largely  by 
conditions  within  the  spur  itself  (Table  11). 

Evidence  from  Chemical  Analysis. — Table  12  records  chemical 
analyses  of  spurs  and  of  bark  from  the  scaffold  limbs  of  bearing  and 
non-bearing  York  trees.  Since  these  data  represent  part  of  the  ma- 
terial which  will  appear  in  a subsequent  bulletin  now  in  prepara- 
tion, no  attempt  is  made  here  to  discuss  their  significance  in  detail, 
though  it  may  be  of  interest  to  note  that  the  bark  even  on  the  scaf- 
fold limbs  of  a tree  in  full  bearing  contains  as  high  percentages 
of  potassium  as  the  spurs,  the  nitrogen  percentage  content  is  only 
haff  of  that  in  the  spurs  and  the  phosphorus  percentage  content 


16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  4 7 


is  at  times  much  less,  but  in  June  of  the  bearing  year  actually 
greater  in  the  bark  than  in  the  spurs. 

Two  points,  however,  should  be  mentioned  for  their  possible 
bearing  here.  Firstly,  the  essential  similarity  in  nitrogen  content 
of  the  bark  and  the  marked  difference  in  this  same  constituent  in 
the  spurs,  as  between  bearing  and  non-bearing  trees  is  at  least 
interesting.  The  second  point  is  the  comparison  of  starch  contents 
which  are  distinctly  different  in  the  bearing  and  non-bearing  years 
and  vary  in  the  bark  in  the  same  manner  as  in  the  spurs.  This 
offers  clear  evidence  that  in  this  respect  a large  part  of  the  aerial 
portion  of  the  tree  is  acting  as  a unit.  The  starch  content  of  the 
bark  may  not  affect  that  of  the  spurs ; both  may  depend  on  the 
same  condition  in  the  tree  as  a whole.  In  other  words,  the  circum- 


Table  12. — The  Chemical  Composition  oe  Bark  and  Spurs  From  Bearing 
and  Non-Bearing  York  Trees. 


Dry 

Reducing  Starch 

Ash 

K 

P 

N 

Weight 

Sugars 

(Per 

(Per 

(Per 

(Per 

(Per 

(Per 

cent 

cent 

cent 

cent 

cent 

cent 

dry 

dry 

dry 

dry 

dry 

dry  weight)  weight)  weight)  weight)  weight) 
weight) 


Trees  in  Bearing 

Bark 


May  22 

43.4 

1.48 

0.11 

7.97 

.621 

.060 

0.58 

June  19 

47.9 

1.32 

0.00 

9.91 

.570 

.145 

0.58 

Sept.  6 

44.9 

1.87 

2.53 

10.76 

.457 

.110 

0.52 

Nov.  20 

44.1 

2.68 

1.58 

12.35 

.498 

.098 

0.69 

Spurs 

May  22 

40.2 

.79 

0.68 

9.73 

.678 

.170 

1.020 

June  19 

44.4 

1.26 

0.00 

6.44 

.554 

.140 

0.916 

Sept.  6 

46.7 

1.21 

2.88 

7.97 

.408 

.155 

1.030 

Nov.  20 

48.4 

2.95 

1.08 

9.05 

.489 

.222 

1.220 

Non-Bearing  Trees 
Bark 

May  27 

42.1 

1.38 

0.72 

11.62 

.601 

.083 

0.56 

June  24 

43.5 

.94 

2.30 

7.85 

.558 

.084 

0.55 

Sept.  20 

48.1 

.90 

3.19 

8.87 

.461 

.113 

0.58 

Dec.  3 

53.8 

4.22 

1.91 

10.28 

.587 

.110 

0.72 

Spurs 

May  27 

45.6 

1.17 

0.92 

8.98 

.593 

.123 

0.773 

June  24 

49.4 

.92 

2.88 

6.67 

.516 

.227 

0.960 

Sept.  20 

53.9 

.79 

2.75 

10.38 

.445 

.202 

0.950 

Dec.  3 

44.7 

2.78 

1.51 

8.85 

.539 

.233 

1.090 

Localization  of  Factors  Determining  Fruit  Bud  Formation  17 


stance  which  prevents  starch  accumulation  in  the  spurs  may  like- 
wise prevent  starch  accumulation  in  the  bark  when  the  majority  of 
spurs  are  bearing  fruit  and  it  might  be  supposed  that  the  large 
amount  of  developing  fruit  which  the  tree  is  bearing  utilizes  all 
the  carbohydrates  that  the  usually  diminished  leaf  area  of  the 
bearing  tree  can  manufacture  and  consequently  diminishes  the 
supply  reaching  the  regions  of  storage.  Since  starch  accumulation 
does  not  occur  in  the  bark  of  these  trees  at  the  time  of  fruit  bud 
differentiation  when  the  tree  is  in  its  bearing  year  it  is  not  sur- 
prising that  starch  accumulation  is  also  absent  in  the  relatively  few 
spurs  that  are  not  bearing  fruit  and  particularly  in  newly  formed 
spurs  on  second  year  wood. 

Hartig9  found  that  previous  to  the  seed  year  in  the  beech  and 
oak  large  amounts  of  starch  were  stored  in  the  medullary  rays  of 
the  wood.  In  non-bearing  years  the  starch  stored  in  the  two 
youngest  annual  ring" was  reduced  to  about  half  after  the  middle  of 
June,  but  the  supply  was  replenished  in  October;  in  the  seed  bear- 
ing year  the  starch  content  of  the  entire  wood  was  reduced  to  a 
minimum,  consuming  the  accumulation  of  eight  years ; further- 
more nearly  all  the  nitrogen  disappeared  from  both  wood  and 
bark,  Hartig  concluded  that  the  food  supply  of  the  buds  was  de- 
rived from  local  accumulations,  that  the  activity  of  the  cambium 
utilized  only  a very  small  amount  of  the  starch  stored  in  the  wood, 
but  that  the  accumulation  of  surplus  reserves  over  a period  of  eight 
years  was  used  in  seed  production.  “In  many  trees,  for  example 
elms  and  fruit  trees,”  he  states,  “a  seed  year  usually  follows  a year 
of  rest  in  which  surpluses  are  accumulated ; in  other  kinds  of  trees 
seed  years  recur  only  after  three,  five  or  even  ten  years.” 

These  findings  are  important  since  they  indicate  that  in  the 
beech  and  oak,  as  well  as  the  alternate  bearing  apple,  the  tree  acts 
more  or  less  as  a unit;  they  suggest  that  the  starch  content  of 
apple  wood  may  be  significant  and  they  imply  a rather  direct  re- 
lation of  the  reserve  starch  in  the  trunk  and  branches  to  the  fruit- 
ful condition  of  the  tree  as  a whole-  If  Hartig’s  surmise  that  the 
starch  accumulations  in  the  trunk  are  used  for  seed  production  be 
correct,  the  question  of  the  passage  of  carbohydrates  up  the  trunk 
must  be  studied  from  a new  point  of  view.  In  the  beech,  for  ex- 
ample, no  significant  upward  translocation  of  carbohydrates  would 
occur  eight  years  out  of  nine.  Very  materially  different  results 
might  be  obtained  from  investigating  trees  in  the  bearing  year 


18 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  47 


than  in  the  off  year  and  a consideration  of  the  condition  of  the  ma- 
terial used  for  experimental  purposes  might  go  far  to  reconcile 
the  conflicting  reports  on  the  upward  translocation  of  carbo- 
hydrates. 

Evidence  from  Diameter  Growth. — The  influence  of  crop  pro- 
duction on  the  older  parts  of  the  tree  is  shown  not  merely  by  the 
chemical  analyses  just  presented,  but  also  by  the  measurements 
of  wood  growth  shown  in  Table  13.  The  greater  width  of  annual 
rings  in  years  when  no  crop  was  borne  cannot  be  attributed  to 
variations  in  climatic  factors  or  in  soil  conditions  since  the  two 
branches  studied  were  taken  from  the  same  tree ; branch  C bore 
fruit  in  even  years  while  branch  D bore  fruit  in  odd  years.  This 


Table  13. — Width  or  Annual  Rings  (in  Millimeters)  on  Two  Branches 

or  a Wealthy  Tree. 


Branch  C 

(bearing  in  even  years) 

Branch  D 

(bearing  in  odd  years) 

Secton  3.3  cm. 
in  diameter 

Section  2.1  cm. 
in  diameter 

Section  3.7  cm. 
in  diameter 

Section  2.3  cm. 
in  diameter 

1917 

1.6 

1.9 

0.5 

1.2 

1918 

0.7 

0.8 

1.2 

0.5 

1919 

1.3 

2.3 

0.3 

0.3 

1920 

0.9 

0.8 

0.4 

1.5 

relationship  has  not  been  studied  sufficiently  to  warrant  an  asser- 
tion of  its  universal  occurrence,  though  the  same  correlation  has 
been  observed  here  in  old  Ben  Davis  spurs  and  McCue10  reports, 
“In  the  alternate  bearing  [apple]  trees,  the  year  of  production  seems 
to  succeed  a year  of  relatively  great  increase  in  trunk  increment.” 

SIGNIFICANCE 

A priori  application  of  these  facts  to  fruit  bud  formation  in  the 
apple  would  be  unjustified  and  is  not  attempted  here.  However, 
enough  evidence  is  available  to  indicate  that  though  conditions  within 
the  spur  are  always  important  and  frequently  decisive,  conditions  in 
the  tree  back  of  the  spur  are  also  important  and  frequently  decisive 
of  the  spur’s  performance.  It  is  shown  here  that  the  spur  or  the 
whole  tree  or  branches  may  be  units  and  that  the  individual  spur  is 
influenced,  sometimes  at  least,  by  the  performance  of  the  other  spurs. 


Localization  of  Factors  Determining  Fruit  Bud  Formation  19 


How  indirect  this  influence  is,  through  effects  on  near  or  on  remote 
portions  of  the  tree,  is  not  yet  determined.  It  is  shown  that  the  col- 
lective performance  of  the  spurs  has  an  influence  on  rather  remote 
parts  of  the  scaffold  limb.  Whether  the  remote  parts  have  an  influ- 
ence on  the  spurs  is  not  yet  shown  definitely.  The  data  presented 
show  that  the  factors  influencing  fruit  bud  differentiation  are  localized 
narrowly  at  times,  widely  at  times.  In  any  case,  careful  investigation 
of  the  factors  determining  fruit  bud  differentiation  should  not  be  con 
fined  to  the  spur  alone. 


REFERENCES 

1.  Lawrence,  J.  The  Clerg3>-- Man's  Recreation,  p.  49.  5th.  ed.  London, 
1717. 

2.  Klebs,  G.  Proc.  Roy.  Soc.  London  82:  547-558.  1910. 

3.  Fisher,  H.  Gartenflora  65:  232-237.  1916. 

4.  Kraus,  E.  J.  and  Kraybill,  H.  R.  Oreg.  Agr.  Exp.  Sta.  Bui.  149.  1918. 

5.  Hooker,  H.  D.  Jr.  Mo.  Agr.  Exp.  Sta.  Res.  Bui.  40.  1920. 

6.  Walster,  H.  L.  Bot.  Gaz.  69:  97-125.  1920. 

7.  Harvey,  E.  M.  and  Murneek,  A.  E.  Oreg.  Agr.  Exp.  Sta.  Bui.  176. 

1921. 

8.  Roberts,  R.  H.  Wis.  Agr.  Exp.  Sta.  Bui.  317.  1920. 

9.  Hartig,  R.  Anatomie  und  Physiologie  der  Pflanzen,  pp.  251-253.  Ber- 
lin, 1891. 

10.  McCue,  C.  A.  Del.  Agr.  Exp.  Sta.  Bui."  126,  1920. 


UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 

AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  48 


INVESTIGATIONS  ON  THE 
HARDENING  PROCESS  IN 
VEGETABLE  PLANTS 

(Publication  Authorized  October  22,  1921) 


COLUMBIA,  MISSOURI 
DECEMBER,  1921 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL, 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY,  P.  E.  BURTON,  H.  J.  BLANTON, 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 

J.  C.  JONES,  Ph.  D.,  LL.  D.,  ACTING  PRESIDENT  OF  THE  UNIVERSITY 


STATION  STAFF 
DECEMBER,  1921 


AGRICULTURAL  CHEMISTRY 

C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  A.  M. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  Sieveking,  B.  S.  in  Agr. 

AGRICULTURAL  ENGINEERING 
J.  C.  Wooley,  B .S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edingf.r,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

E.  F.  Hopkins,  Ph.  D. 

DAIRY  HUSBANDRY 
C.  Ragsdale.  B.  S.  in  j\gr. 

. W.  Swett,  A.  M. 

M.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride, 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  A.  M. 

O.  W.  Letson,  B.  S.  in  Agr. 

B.  M.  King,  B.  S.  in  Agr. 

A.  C.  Hill,  B.  S.  in  Agr. 

Miss  Bertha  C.  Hite,  A.  B.1 
Miss  Pearl  Drummond,  A.  A.1 


RURAL  LIFE 

O.  R.  Johnson,  A.  M. 

S.  D.  Gromer,  A.  M. 

E.  L.  Morgan,  A.  M. 

Ben  H.  Frame,  B.  S.  in  Agr. 


HORTICULTURE 

V.  R.  Gardner,  M.  S.  A. 

H.  D.  Hooker,  Jr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  M.  S. 

F.  C.  Bradford,  M.  S. 

H.  G.  Swartwout,  B S.  in  Agr. 

POULTRY  HUSBANDRY 

H.  L.  Kempster,  B.  S. 

Earl  W.  Henderson 


SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M. 

W.  A.  Albrecht,  Ph.  D. 

F.  L.  DulEy,  A.  M.2 
R.  R.  HudElson.  A.  M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr. 
Richard  Bradfield,  A.  B. 

O.  B.  Price,  B.  S.  in  Agr. 

VETERINARY  SCIENCE 

J.  W.  Connaway,  D.  V.  S.,  M.  D. 
L.  S.  Backus,  D.  V.  M. 

O.  S.  Crtsler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 


OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Sercretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Teffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Jane  Frodsham,  Librarian 
E.  E.  Brown,  Business  Manager 


Hn  service  of  U.  S.  Department  of  Agriculture,  Seed  Testing  Laboratory. 
2On  leave  of  absence. 


TABLE  OF  CONTENTS. 


Page 

Introduction  5 

Review  of  Literature  5 

The  physical  process  of  freezing  in  plants  5 

Nature  of  the  killing  of  plant  tissue  by  cold  10 

Result  of  water  loss-desiccation  11 

Injury  to  Plasma  Membrane  by  water  withdrawal  12 

Protein  precipitation  through  “salting  out”  13 

Protein  precipitation  by  increase  in  acidity  13 

Relation  of  water  withdrawal  from  the  cells  to  killing  by  cold  14 

Factors  influencing  the  water-retaining  power  of  cells 15 

Osmotic  concentration  and  water-retaining  power  15 

Imbibition  and  water-retaining  power  17 

Relation  of  factors  influencing  water-loss  by  the  plant  as  a whole,  to 

hardiness  22 

Statement  of  the  problem  25 

Experimental  work  20 

Materials  used  20 

Methods  of  hardening  plants  26 

Effect  of  hardening  treatments  on  plants  28 

Morphological  differences  in  hardened  plants  29 

Effect  of  hardening  treatments  on  rate  of  growth  30 

Effect  of  hardening  treatments  on  percentage  of  dry  matter 31 

Effect  of  hardening  treatments  on  depression  of  freezing  point 32 

Effect  of  hardening  treatment  on  ice  formation  in  plants 37 

Method  of  measuring  the  amount  of  water  freezing  in  plant  tissues  . . 41 
Effect  of  temperature  on  amount  of  water  freezing  in  hardened  and 

non-hardened  cabbage  leaves  42 

Changes  in  amount  of  freezable  water  during  the  hardening  process  . 45 

Influence  of  time  of  day  on  percentage  of  water  frozen  47 

Effect  of  watering  plants  with  salt  solutions  on  amount  of  easily 

frozen  water  in  the  leaves  48 

Relation  of  amount  of  freezable  water  to  percentage  of  dry  matter 

and  freezing  point  depression  in  garden  plants  53 

Rate  of  water-loss  by  transpiration  in  hardened  and  tender  cabbage  ....  55 

Rate  of  dehydration  in  hardened  and  tender  plants  58 

Changes  in  Carbohydrates  on  hardening  of  plants  60 

Formation  of  sugar  by  low  temperature  60 

Relation  of  sugar  content  to  cold  resistance 61 

Methods  of  analysis — carbohydrates  64 

Nature  of  water-retaining  power  in  plants  66 

Relation  of  pentosan  content  to  cellular  water-retaining  power 68 

Pentosan  content  in  the  hardening  process  in  vegetable  plants 70 

3 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Method  of  pentosan  analysis  70 

Pentosan  content  in  garden  plants  72 

Pentosan  content  in  plants  watered  with  salt  solutions 73 

Rate  of  increase  in  pentosan  content  73 

Relation  of  hot  water  soluble  pentosans  to  the  hardening  process 75 

Factors  influencing  the  imbibitional  capacity  of  plant  colloids 77 

Acidity  77 

Salts  and  Sugars  80 

Summary 81 

Conclusions  S3 

Applications  81 

Acknowledgments  85 

Bibliography  85 

Plates  91 


INVESTIGATIONS  ON  THE  HARDENING 
PROCESS  IN  VEGETABLE  PLANTS 

J.  T.  Rosa,  Jr. 

This  study  was  undertaken  as  one  phase  of  a project  on  the 
transplanting  of  vegetable  plants.  The  hardening  process,  whereby 
vegetable  plants  are  made  more  resistant  to  cold  and  better  able  to 
withstand  the  hardships  of  transplanting  from  greenhouse  or  hotbed 
to  the  open  field,  is  of  great  importance  in  the  practice  of  growing 
certain  vegetables  which  are  customarily  transplanted.  In  the  pro- 
duction of  early  crops,  hardiness  also  is  especially  important  be- 
cause of  the  low  temperatures  to  which  transplanted  plants  are  ex- 
posed upon  their  removal  to  the  field  in  early  spring. 

Furthermore,  since  the  hardening  process  in  vegetable  plants 
results  in  a condition  of  acquired  hardiness,  developed  rather  quick- 
ly by  subjecting  plants  to  certain  treatments,  experiments  with  such 
material  throw  considerable  light  on  the  general  problem  of  cold  re- 
sistance in  plants.  This  question,  in  connection  with  that  of  the 
nature  of  the  process  of  killing  of  plants  by  low  temperature,  has 
received  the  attention  of  numerous  investigators  during  the  past 
one  hundred  years.  Though  much  information  has  been  accumu- 
lated, the  whole  problem  is  in  a somewhat  undefined  state.  It  is 
the  purpose  of  this  paper  to  propose  a theory  comprehensive  enough 
to  explain  satisfactorily  the  known  facts  as  to  the  cold-resistance  of 
living  plants  and  to  present  data  on  the  nature  of  the  response  of 
plant-tissues  to  treatments  which  result  in  increased  hardiness.  The 
injurious  effects  of  temperature  slightly  above  the  freezing  point  on 
the  growth  of  plants  are  not  dealt  with  in  this  paper. 

REVIEW  OF  LITERATURE. 

The  Physical  Process  of  Freezing  in  Plants. — An  early  theory 
as  to  killing  of  plants  by  cold,  advanced  by  Duhamel  and  Buffon27* 
in  1737,  held  that  death  was  due  to  the  rupture  of  the  tissues, 
bursting  of  the  plant  cells,  by  the  expansion  of  ice  crystals  form- 
ing within  the  cells  upon  freezing. 

•This  and  subsequent  superscript  numerals  refer  to  literature  cited  in  the  Bibliography. 
NOTE. — Also  submitted  to  the  Faculty  of  the  Graduate  School  of  the  University  of  Mis- 
souri as  a thesis  in  partial  fulfilment  of  the  requirements  for  the  degree  of  Doctor  of  Philos- 
ophy. 

5 


6 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Geoppert32  in  1829,  found  that  ice  formation  upon  freezing  of 
plant  tissue  was  not  confined  to  the  interior  of  the  cells  and  con- 
cluded that  the  killing  of  plants  by  cold  was  not  due  to  cell  rupture. 
A few  years  later,  Morren73  substantiated  Geoppert ’s  conclusion  in 
that  he  found  no  organ  of  the  plant  torn  by  freezing.  He  consid- 
ered that  injury  from  freezing  was  due  mostly, to  the  separation  of 
air  from  the  plant  sap.  In  1860,  Sachs105  using  improved  technique, 
observed  that  in  the  process  of  freezing,  water  was  withdrawn  from 
the  cell  and  ice-crystals  formed  for  the  most  part  in  intercellular 
spaces. 

In  1860,  Nageli89  showed  by  calculation,  that  the  expansion 
caused  by  freezing  all  the  water  in  the  cell,  would  not  be  sufficient 
to  cause  a rupture  of  the  cell-wall.  Prillieux"  in  1869,  found  that 
water  was  extruded  from  the  cells  upon  freezing.  Muller-Thurgau74 
found  that  ice  formed  within  the  cell  to  some  extent,  when  the 
lowering  of  the  temperature  was  very  rapid,  but  in  case  of  gradual 
cooling  to  the  point  of  ice  formation  as  in  nature,  the  crystals  were 
found  exclusively  in  the  intercellular  spaces.  Wiegand28  noted 
similar  results  upon  freezing  Spirogyra  and  Nitella.  Thus  the  find- 
ing of  ice  crystals  within  the  cells  by  earlier  investigators,  who  froze 
the  plant  tissue  very  quickly,  is  explained.  Cavallero19  confirmed 
the  work  of  the  German  writers,  as  he  found  that  cell  rupture  in 
winter  was  very  rare,  the  cells  themselves  never  freezing,  though 
ice  formation  occurred  in  the  intercellular  spaces  of  both  hardy  and 
tender  plants. 

Geoppert32  noted  that  plants  which  were  frozen  to  death  lost 
water  rapidly  upon  thawing.  Sachs106  observed  that  upon  thawing, 
water  remained  in  the  intercellular  spaces  until  reabsorbed  by  the 
cells  or  lost  by  evaporation.  Under  certain  conditions  considerable 
time  elapsed  before  the  water  was  reabsorbed  and  the  protoplast  re- 
gained its  turgid  condition.  Prillieux100  describes  experiments  on 
freezing  pieces  of  potato  and  beet,  showing  that  water  was  lost 
from  these  tissues  upon  thawing.  He  recognized  also  that  water  was 
lost  from  the  tissues  while  still  frozen,  by  evaporation  from  the  sur- 
face of  the  ice  crystals. 

Prunet101  found  that  moisture  is  lost  by  evaporation  from  the 
surface  of  the  leaf  on  thawing,  rather  than  by  normal  transpiration 
through  the  stomata. 

Abbe1  stated  that  as  plant  tissues  were  cooled,  water  exuded 
from  the  cells  into  the  intercellular  spaces,  and  after  sufficient  under- 


The  Hardening  Process  in  Vegetable  Plants. 


7 


cooling,  this  water  froze.  The  concentrated  sap  left  within  the  cell 
did  not  freeze  until  cooled  still  lower. 

If  the  water  is  withdrawn  from  the  cell  before  freezing  in  the 
intercellular  spaces,  it  is  important  to  find  how  this  withdrawal 
takes  place.  Wiegand131  offered  two  theories  to  account  for  cellular 
water  loss  upon  freezing,  ‘ ‘ extrusion  ’ ’ and  ‘ ‘ attraction.  ’ ’ 

Extrusion. — This  hypothesis  is  that  the  cell  actively  gives  up 
water  at  low  temperature  by  contraction  and  squeezing.  Greeley34 
showed  that  cooling  to  near  0°C.  caused  Stent  or  to  contract  and 
become  cyst-like.  Under  the  same  conditions  Spirogyra  became  much 
plasmolyzed.  Livingston  showed  that  when  mounted  in  oil,  this 
plasmolysis  was  accompanied  by  extrusion  of  droplets  of  water. 
Wiegand  thought  that  the  most  probable  explanation  of  this  method 
of  water  loss  from  the  cell  was  by  change  in  permeability  of  the 
protoplast  to  the  sap  solute.  A recent  report  by  Pantanelli96  sup- 
ports this  idea.  In  experiments  with  the  pericarp  of  the  mandarin 
cooled  almost  to  the  freezing  point  of  this  material  (-6°C.)  he 
observed  a progressive  increase  in  cellular  permeability,  as  shown 
by  rapid  loss  of  water  and  exomosis  of  substances  from  the  tissue. 
Osterhout93  has  shown  that  freezing  as  well  as  treatment  by  various 
anesthetics,  greatly  increases  cellular  permeability. 

Attraction. — Wiegand139  considered  his  so-called  attraction  the- 
ory as  the  more  probable  explanation  of  water  withdrawal  from 
the  cell.  Thus  in  ordinary  plant  tissue  Wiegand  pictured  the  fol- 
lowing arrangement: 

(1)  A film  of  pure,  or  nearly  pure,  water  adhering  to  the  outer 
surface  of  the  cell  wall,  bordering  on  the  intercellular  spaces. 

(2)  The  inert  cell-wall  cellulose  material  filled  with  water  of 
imbibition,  which  is  continuous  with  that  of  the  protoplast. 

(3)  A more  or  less  narrow  strip  of  protoplasm  adhering  close- 
ly to  the  inner  surface  of  the  cell  wall  and  containing  water  of  im- 
bibition, continuous  with  that  of  the  vacuole. 

(4)  The  vacuole,  containing  an  aqueous  solution  of  salts,  sug- 
ars and  other  substances. 

Normally  this  system  is  in  equilibrium.  According  to  Wiegand, 
upon  lowering  the  temperature  below  the  freezing  point,  the  film 
of  pure  water  on  the  outer  surface  of  the  cell  walls  freezes  first. 
The  tendency  will  then  be  to  restore  equilibrium  by  drawing  water 
from  the  interior  of  the  cell  to  replace  the  surface  film.  This  water 
will  be  drawn  first  from  the  cell  wall,  which  in  turn  will  draw  on  the 


8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


protoplast,  which  in  turn  will  draw  on  the  sap  in  the  vacuole.  The 
water  of  the  vacuole  is  held  by  the  force  of  solution  alone,  whereas 
the  cell  wall  and  protoplasm  hold  water  by  the  stronger  force  of 
imbibition.  If  the  temperature  remains  constant,  this  readjustment 
will  continue  until  the  force  of  crystallization  is  equalled  by  the  in- 
creased force  with  which  the  remaining  water  is  held  within  the  cell. 
After  equilibrium  is  established  between  the  forces  of  crystallization 
and  the  water-retaining  power  of  the  cell,  at  any  given  temperature, 
no  more  water  freezes  unless  the  temperature  is  lowered  further, 
thereby  increasing  the  force  of  crystallization.  However,  since  the 
force  with  which  the  remaining  water  is  held  increases  rapidly  with 
the  progressive  loss  of  water,  Wiegand  predicted  that  the  amount  of 
water  frozen  at  each  successive  degree  for  which  the  temperature 
is  lowered  would  be  smaller  and  smaller.  This  was  shown  to  be  ap- 
proximately true  by  the  experiments  of  Muller-Thurgau74  with  ap- 
ples, and  the  work  of  McCool  and  Millar80  with  green  plants  sug- 
gests the  same  conclusion.  Bouyoucos11  working  with  soils,  found 
that  little  more  water  was  frozen  at  -78 °C.  than  at  -6°C. 

The  foregoing  hypothesis  as  to  the  conditions  under  which  ice 
is  formed  in  living  plant  tissue  has  been  substantiated  by  work  of 

Effect  of  Glucose  Solutions  on  Cold  Resistance  in  Sections  of  Red  Cab- 
bage Leaves. 


Temperature 

Concentration  of  Solution. 

2M 

M 

M/2 

M/4 

M/8 

M/16 

Water 

- 5.2°C. 

all 

living 

Vz  cells 
alive 

- 7.8°C. 

all 

living 

14  cells 
alive 

single 

cells 

alive 

all 

dead 

•ii.rc. 

all 

living 

y2  cells 
alive 

single 

cells 

alive 

all 

dead 

-17. 3°C. 

all 

living 

y2  cells 
alive 

single 

cells 

alive 

all 

dead 

-22.0°C. 

all 

living 

single 

cells 

alive 

all 

dead 

-32°C. 

Vo 

cells  | 
alive  1 

single 

cells 

alive 

all 

dead 

The  Hardening  Process  in  Vegetable  Plants. 


9 


Maximow.66  In  extensive  experiments  with  red  cabbage  and  Trades- 
cantia  discolor  he  found  a marked  ‘ ‘ protective  ” action  when  sec- 
tions were  frozen  in  solutions  of  salts,  sugars,  and  other  organic  ma- 
terials, provided  the  substance  used  was  not  toxic  and  its  eutectic 
point  did  not  lie  too  near  the  freezing  point.  Although  the  con- 
ditions of  Maximov/  ’s  experiments  cannot  be  duplicated  in  nature, 
his  results  are  of  interest.  The  following  table,  taken  from  Maxi- 
mow’s work,  is  typical  of  the  results  he  secured. 

Evidently  red  cabbage  cells,  which  ordinarily  are  killed  at  a little 
below  -5°C.,  survive  a temperature  as  low  as  -32°C.  in  2-mol.  glucose 
solution.  Maximow  concluded  that  this  apparent  protective  action 
of  the  solution  could  not  be  explained  by  the  depression  of  the  freez- 
ing point,  since  the  resistance  to  cold  always  increased  with  the 
strength  of  the  solution  much  more  rapidly  than  this  depression. 
The  degree  of  protection  was  found  however,  to  be  closely  related  to 
the  eutectic  point  of  the  solution,  substances  having  a high  eutectic 
point  showing  no  protective  effect.  Isotonic  solutions  of  different 
substances  with  low  eutectic  points  possessed  nearly  the  same  degree 
of  protective  action.  Maximow  found  no  relation  between  the  rate 
of  penetration  of  the  protective  substance  and  the  degree  of  protec- 
tion afforded,  and  that  just  as  much  protective  action  was  exerted 
by  the  various  solutions  when  sections  were  immersed  in  them  and 
frozen  immediately,  as  when  the  tissue  had  been  soaked  several  hours 
in  the  solution  before  freezing.  (Hence  there  could  have  been  no 
effect  on  cell  sap  concentration  or  in  preventing  precipitation  of  the 
cell  proteins.) 

If  we  consider  Maximow’s  work  in  connection  with  Wiegand’s 
hypothesis  of  freezing,  we  have  a condition  differing  from  the  usual, 
in  that  the  film  of  pure  water  on  the  outer  surface  of  the  cell  wall 
is  replaced  by  a more  or  less  concentrated  solution.  In  the  first  place, 
this  would  lower  the  initial  freezing  point  somewhat.  More  im- 
portant still,  the  fact  that  the  cell  is  surrounded  by  a more  or  less 
concentrated  solution  should  mean  that  in  the  process  of  water  with- 
drawal and  ice  formation  at  any  given  temperature,  a state  of  equi- 
librium between  the  ice-ciystal  and  the  cell  system  would  be  reached 
sooner  than  in  the  case  of  cells  not  surrounded  by  such  solutions,  if 
the  ‘‘attraction”  theory  of  water  loss  as  advanced  by  Wiegand  be  ac- 
cepted. Somewhat  less  water  would  be  frozen  at  a given  temperature 
in  the  cells  of  tissue  immersed  in  salt  or  sugar  solution.  If  the  amount 
of  water  frozen  per  degree  of  temperature  lowering  becomes  smaller 
and  smaller,  it  would  be  necessary  for  a “protective”  solution  to 


10 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  43 


effect  a very  small  reduction  in  the  amount  of  water  freezing  at  the 
lower  temperatures  to  enable  the  cell  to  stand  cooling  several  degrees 
below  the  usual  death-point.  Recent  work  by  Yass  (122)  on  bacteria 
leads  to  the  same  conclusion.  He  found  a distinct  protective  action 
exerted  by  glycerine  and  glucose  solutions  on  freezing  bacteria,  as 
shown  in  the  following  table. 

Vass’  Results  on  Freezing  of  Bacteria  at  -5°C. 


Strength  of  solution 

Percent 

of  bacteria  killed 

In  glycerine 

In 

glucose 

0.00  (water) 

96 

— 

0.01% 

92 

98 

0.05% 

87 

95 

0.1 

41 

89 

0.5 

45 

74 

1.0 

0 

58 

5.0 

0 

35 

10.0 

0 

4 

Vass  concluded,  in  agreement  with  Maximow,  that  the  protective 
action  of  these  solutions  was  due  to  their  power  to  keep  a film  of  un- 
frozen water  in  contact  with  the  outer  layer  of  the  protoplast,  the 
plasma  membrane. 

Nature  of  the  killing  of  plant-tissue  by  cold. — From  the  fore- 
going review,  the  evidence  appears  conclusive  that  cell  rupture  cannot 
be  the  cause  of  killing  of  plants  by  cold,  but  that  water-loss  from  the 
cells  by  ice  formation  in  the  intercellular  spaces  is  an  invariable 
accompaniment  in  such  killing.  According  to  Muller-Thurgau,76  Mo- 
lisch71  and  others,  death  cannot  be  due  directly  to  absolute  cold, 
and  there  is  little  if  any  evidence  of  death  due  to  shock  or  other  re- 
action attributable  to  ‘ ‘ cold-rigor.  ” Thus,  both  Muller-Thurgau76 
and  Voightlander123  showed  that  plant  tissues  could  be  undercooled 
several  degrees  below  the  freezing  point  without  injury  as  long  as  ice 
formation  did  not  take  place.  Wright  and  Taylor122  have  recent- 
ly shown  that  potatoes  can  be  cooled  several  degrees  below  their  freez- 
ing point  and  warmed  up  again  without  injury,  provided  no  ice 
formation  took  place.  However,  jarring  undercooled  potatoes  caused 
ice-formation  to  take  place  and  resulted  in  typical  frost  injury. 

Chandler20  found  evidence  that  tender  plants  exposed  to  tem- 
perature slightly  below  freezing  when  the  surface  of  the  leaves  was 
wet,  killed  to  a greater  extent  than  if  the  leaves  were  dry.  This 
result  is  explained  by  Harvey's  “injection"  theory,  according  to  which 
undercooled  tissues  are  caused  to  freeze  in  spots  where  droplets  of 
free  water  on  the  surface  crystallize  and  inoculate  the  tissue  just 
beneath  with  the  growing  crystals. 


The  Hardening  Process  in  Vegetable  Plants. 


11 


These  facts  strengthen  the  view  that  killing  by  cold  depends  on 
ice  formation,  rather  than  on  the  effect  of  low  temperature  in  itself. 
Just  how  death  is  caused  by  the  freezing  process  is  a question  of 
interest.  Four  distinct  theories  have  been  advanced. 

( a ) Direct  result  of  water  loss — “ desiccation — Miiller-Thur- 
gau74  believed  that  death  was  the  direct  result  of  the  water  loss, 
that  is,  death  ensues  when  so  many  molecules  of  water  are  withdrawn 
from  the  protoplast  that  its  living  structure  is  permanently  destroyed. 
Wiegand131  concurred  in  this  hypothesis,  with  the  additional  sugges- 
tion that  “probably  every  cell  has  its  critical  point,  beyond  which 
water  withdrawal  causes  death.”  Cavallero19  attributed  killing  to 
the  wilting  upon  thawing,  due  to  rapid  evaporation  of  melting  ice 
in  the  intercellular  spaces.  He  mentioned  an  opinion  generally  held 
by  practical  gardeners.,  that  under  conditions  favoring  slow  thawing 
or  slow  evaporation,  such  as  shade  or  moisture,  severe  injury  to  the 
plant  might  be  prevented  by  the  re-entry  of  water  into  the  cells. 
However,  Miiller-Thurgau74  and  later  Molisch71  found  no  difference 
in  extent  of  killing,  between  rapid  and  slow  thawing.  Chandler20 
also  concluded  from  a considerable  number  of  experiments  that  the 
rate  of  thawing  generally  had  no  influence  on  death  from  freezing. 
We  should  distinguish  here  between  the  loss  of  water  from  the  cell 
and  its  loss  from  the  plant  as  a whole.  If  the  cells  are  killed  directly 
by  loss  of  water  on  freezing,  or  if  they  are  killed  by  changes  taking 
place  as  a result  of  this  water  loss,  then  the  rate  of  thawing  would 
have  no  effect  on  the  killing.  However  plants  capable  of  standing 
some  ice  formation  within  their  tissues,  would  take  back  more  of  this 
water  if  thawed  slowly,  whereas  they  might  lose  the  most  of  it  if 
thawed  rapidly.  This  explains  two  things,  the  wilted  condition  often 
observed  in  frozen  plants  upon  thawing,  and  the  cumulative  effect  of 
successive  freezing  and  thawing,  whereby  a fraction  of  the  plant’s 
water  content  is  permanently  lost  by  the  plant  on  each  thawing. 

Nelson90  thought  that  rapid  loss  of  moisture  was  the  principal 
cause  of  winter-killing  of  shrubs  in  high,  dry  sections.  Kylin56  in  a 
recent  study  of  the  cold  resistance  of  marine  algae,  concluded  that 
death  from  cold  was  conditional  upon  actual  formation  of  ice  and 
that  such  death  was  primarily  due  to  withdrawal  of  water  from  the 
cell.  Matruchot  and  Molliard04  described  the  successive  changes  in 
arrangement  of  the  chromatin  strands  of  the  nuclei  in  leaf  cells  of 
the  snowdrop  subjected  to  freezing  temperatures.  They  stated  that 
water  was  withdrawn  from  the  protoplast  and  nuclear  material  of  the 
cell,  and  that  this  continued  if  the  temperature  was  sufficiently  low, 


12 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


until  these  portions  of  the  cells  contained  less  water  than  the  mini- 
mum necessary  to  vitality.  They63  also  subjected  plant  tissue  to 
freezing,  to  drying  and  to  the  action  of  solutions  of  high  osmotic 
concentrations.  They  observed  a marked  parallelism  in  the  effects  of 
these  treatments,  hence  they  concurred  with  Molisch  and  Miiller- 
Thurgau  in  that  death  of  the  cell  was  due  to  rapid  loss  of  water. 
Adams3  working  with  moist  seeds,  observed  that  in  freezing,  water 
was  drawn  from  the  cells  and  solidified  in  the  intercellular  spaces 
and  if  the  freezing  did  not  go  too  far,  upon  melting  the  water  was 
reabsorbed  slowly,  without  injury  having  been  done  to  the  cells. 

Wiegand129  made  extensive  observations  on  the  freezing  of  leaves 
and  buds.  In  buds  in  winter:  “Ice  was  always  found  in  broad, 
prismatic  crystals  arranged  perpendicularly  to  the  excreting  surface 
and  usually  formed  a single  continuous  layer  throughout  the  meso- 
phyl  of  the  scale  or  leaf,  to  accommodate  which  the  cells  were  often 
separated  a considerable  distance.  The  cells  near  the  ice  mass  having 
lost  their  water,  were  in  a state  of  collapse,  but  upon  thawing  they 
reabsorbed  the  water  and  resumed  their  normal  condition.”  This 
was  also  true  of  evergreen  leaves,  in  which  he  observed  that  ice 
crystals  first  lined  the  spaces  of  the  spongy  parenchyma,  later  filling 
these  spaces  and,  in  leaves  of  high  water-content,  the  crystals  fused 
into  a sheet  of  ice  completely  separating  the  upper  and  lower  por- 
tions. 

In  observations  on  thawing  of  frozen  leaf  sections,  Wiegand  no- 
ticed in  hardy  tissues,  not  killed  by  the  freezing,  that  upon  thawing 
the  water  was  drawn  back  into  the  cells,  but  in  tender  tissues  killed 
by  the  freezing  process,  water  was  not  drawn  back  into  the  cells  to 
any  extent.  Pantanelli94  concluded  that  the  suffrance  of  each  cell 
is  directly  proportional  to  the  outgo  of  water  during  cooling.  He  also 
attaches  great  importance  to  the  condition  of  the  roots  with  reference 
to  ready  water  absorption  in  determining  whether  or  not  plant  re- 
covers from  freezing. 

(b)  Injury  to  the  plasma  membrane  by  water  withdrawal. — 
Maximow66  concluded  as  a result  of  an  extensive  series  of  experiments, 
wherein  sections  of  plants  were  frozen  in  solutions  of  various  salts 
and  inorganic  materials,  that  killing  by  cold  is  not  due  to  low  tem- 
perature as  such,  but  to  physico-chemical  changes  set  up  in  the  col- 
loids of  the  plasma  membrane  during  ice  formation  therein.  This  is 
really  a modification  of  Miiller-Thurgau ’s  theory,  limiting  the  in- 
jurious effects  of  water-loss  to  the  outer  layers  of  the  protoplast. 

Chandler20  also  concluded  from  his  exhaustive  researches  that 


The  Hardening  Process  in  Vegetable  Plants. 


13 


“killing  from  cold  is  more  likely  a mechanical  injury  due  to  with- 
drawal of  water  from  the  protoplasmic  membrane  than  an  injury 
resulting  from  a precipitation  of  proteins.  ” 

(c)  Protein  precipitation  through  “ salting  out.” — Gorke36  con- 
cluded that  killing  was  due  to  irreversible  precipitation  of  the  pro- 
teins of  the  cell.  He  accounts  for  this  precipitation  by  the  greater 
concentration  of  the  salts  in  the  sap  as  water  is  withdrawn  from  the 
cell  by  formation  of  ice,  since  certain  proteins  are  precipitated  in 
strong  salt  solutions.  He  found  that  approximately  l/3  of  the  pro- 
teins were  precipitated  in  frozen  cereal  plants.  Gorke  found  also 
that  hardiness  of  certain  plants  bears  some  relation  to  the  ease  with 
which  their  proteins  were  precipitated.  In  the  tender  begonia  he 
obtained  protein  precipitation  at  -3°C.,  in  winter  rye  at  -15 °C.  and 
in  pine  needles  at  -40 °C.  Schaffnit110  also  concluded  that  protein 
precipitation  was  the  cause  of  death.  He  found  that  the  proteins 
of  rye  plants  grown  in  the  open  at  low  temperature  were  not  as  easily 
precipitated  upon  freezing  as  those  of  tender  greenhouse  plants. 
The  effect  of  low  temperatures  on  the  hardiness  of  plants  grown  in 
the  open  was  ascribed  to  a transition  from  less  stable  to  more  stable 
forms  of  the  proteins  by  splitting.  He  found  that  he  could  prevent 
the  precipitation  of  proteins  from  the  sap  of  tender  greenhouse  plants 
by  addition  of  sugar,  to  which  he  ascribed  a protective  action  against 
protein  precipitation  and  consequently  against  injury  of  the  plant 
from  cold,  although  it  was  not  proven  that  these  two  are  always 
related. 

Chandler20  was  rather  disinclined  to  accept  the  idea  of  killing 
by  “salting  out”  of  proteins.  He  found  that  the  hardiness  of  plants 
was  increased  by  growing  them  in  salt  solutions,  such  as  zinc  sulphate, 
which  is  an  excellent  protein-coagulating  agent.  However,  Chand- 
ler’s work  on  this  point  cannot  be  held  to  disprove  the  protein-pre- 
cipitation idea,  since  he  showed  no  evidence  that  the  protein-precipi- 
tating salts  were  taken  up  by  the  plant,  or  if  they  were  taken  up, 
that  they  existed  in  the  plant  in  a form  which  would  precipitate 
proteins  upon  concentration.  However,  the  fact  that  Chandler  did 
not  find  appreciable  protein  precipitation  on  freezing  the  extracted 
sap  of  apple  twigs  indicates  that  killing  may  not  always  be  accom- 
panied by  protein  precipitation,  although  his  technique  on  this  point 
may  be  open  to  question. 

(d)  Protein  precipitation  by  increase  in  acidity. — Changes  in 
color  of  plant  sap  due  to  change  in  reaction  upon  freezing  are  well 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


known.  Gorke35  noted  an  increase  of  acidity  in  sap  upon  freezing. 
He  believed  this  was  a factor  in  the  precipitation  of  the  plant  pro- 
teins, since  the  acidity  of  the  medium  is  important  in  determining 
the  state  of  such  colloidal  materials. 

Harvey,42  in  a recent  paper  dealing  with  cold  injury  to  cabbage 
plants,  extended  this  theory.  He  found  definite  evidence  of  increased 
acidity  as  a result  of  freezing  cabbage  plant  juice,  by  measuring  the 
hydrogen-ion  concentration  before,  during  and  after  freezing  to 
definite  temperatures.  He  noted  protein  precipitation  when  the 
actual  acidity  was  increased  from  Ph  5.65  to  Ph  5.26.  It  is 
especially  interesting  to  note  that  Harvey  found  a similar  increase 
in  the  acidity  of  juice  expressed  from  leaves  exposed  to  wilting, 
though  he  does  not  state  if  the  leaves  were  wilted  beyond  recovery. 
Harvey  demonstrated  that  if  phosphoric  acid  was  added  to  the  ex- 
pressed sap  until  the  Hydrogen-ion  concentration  was  increased  as 
much  as  it  would  have  been  by  freezing,  a precipitation  of  the  protein 
occurred,  thus  implying  that  the  parallel  effect  of  water  loss  by  wilt- 
ing or  by  freezing  and  addition  of  acid,  was  protein  precipitation  and 
death. 

Harvey  repeated  Gorke ’s  experiment  on  the  precipitation  of  pro- 
tein from  expressed  sap  by  freezing.  Samples  of  juice  were  taken 
from  hardened  and  not  hardened  cabbage  plants  and  frozen  to  -4°C., 
a temperature  which  would  kill  the  non-hardened,  but  not  the  har- 
dened plants.  It  was  found  that  9.4  percent  of  the  protein  in  the 
juice  of  the  hardened  plants  was  precipitated  and  31.2  percent  in 
the  tender  plants.  Repeating  the  experiment  and  adding  sufficient 
acid  to  change  the  reaction  of  the  juice  the  same  amount  as  it  would 
be  changed  by  freezing  to  -3°C.  he  found  that  11  percent  of  the 
protein  was  precipitated  in  the  juice  of  hardened,  and  44  percent 
in  tender  plants.  He  also  made  complete  analyses  of  hardened  and 
tender  cabbage  plants,  finding  that  of  the  water-soluble  fraction  of 
nitrogen  about  35  percent  was  amino-nitrogen  in  hardened  plants, 
and  only  17  percent  in  tender  plants,  having  about  the  same  amount 
of  water-soluble  nitrogen.  Harvey  thought  this  increase  in  amino- 
nitrogen  to  be  a very  significant  result  of  the  hardening  process, 
though  he  said  it  was  not  necessary  that  complete  cheavage  of  the 
proteins  to  the  amino  acids  should  occur,  to  prevent  their  precipita- 
tion on  freezing. 

Relation  of  water- withdrawal  from  the  cells  to  killing  by 
cold. — No  matter  which  agency  is  chiefly  operative  in  the  actual 
freezing  and  killing  process,  they  all  depend  on  the  withdrawal  of 


The  Hardening  Process  in  Vegetable  Plants. 


15 


water  from  the  cell.  Irreversible  coagulation  of  colloids,  such  as 
protoplasm,  is  itself  essentially  a dehydration  process.  It  is,  then, 
by  means  of  factors  affecting  water-withdrawal  from  the  cell  by  ice- 
formation  that  the  differential  killing  of  plant  tissues  by  low  tem- 
peratures may  be  explained. 

Schaffnit110  classified  plants  in  three  groups,  according  to  their 
cold-resistance  and  ability  to  withstand  desiccation. 

1.  Plants  for  which  water  is  absolutely  essential.  This  we  take 
to  include  such  plants  as  tomatoes,  which  are  killed  once  extensive 
ice  formation  actually  takes  place. 

2.  Plants  which  withstand  a certain  degree  of  desiccation. 
These  would  be  such  plants  as  the  cabbage  which  can  survive  a cer- 
tain amount  of  ice-formation  in  the  tissues  without  injury.  It  is  this 
group  with  which  we  are  mostly  concerned  in  discussions  of  harden- 
ing or  cold-resistance. 

3.  Those  which  withstand  complete  drying — seeds,  spores,  etc. 

This  classification  can  be  taken  to  include  all  plants,  except 

those  which  are  killed  by  cold  above  the  freezing  point.  Such  killing 
is  probably  due  to  inability  to  carry  on  their  normal  metabolic  func- 
tions at  low  temperatures,  as  suggested  by  Molisch,  rather  than  to 
direct  effect  of  cold. 

Relationship  to  cold  resistance  of  factors  influencing  the  water- 
retaining  power  of  cells. — If  the  killing  of  plant  tissue  by  cold  is 
primarily  due  to  water-withdrawal  from  the  cells  beyond  a certain 
minimum  point,  then  the  difference  between  hardy  and  tender  tissues 
may  be  ascribed  largely  to  the  relative  water-retaining  power  of  the 
cells  in  the  two  types  of  tissue. 

There  are  two  main  forces  concerned  in  the  water- retaining 
power  of  plant  cells.  (1)  Osmotic  concentration,  due  to  sap  solutes 
in  the  vacuole,  and  (2)  Imbibition,  a force  exerted  by  some  con- 
stituents of  the  cell  wall,  nucleus,  plastids,  and  especially  by  the 
colloidal  cytoplasm.  The  importance  of  either  of  these  forces  in  the 
water-retaining  power  of  cells  may  be  influenced  by  various  factors. 

Osmotic  concentration  and  water -retaining  power. — Since  the 
freezing  point  of  a solution  is  lowered  in  proportion  to  its  molecular 
concentration,  several  workers  have  sought  a correlation  between  coM 
resistance  and  the  molecular  concentration  of  the  sap  as  measured 
by  the  depression  of  the  freezing  point. 

Lindley01  in  reviewing  the  work  of  Morron  and  others  in  1852, 


16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


was  probably  the  first  writer  to  connect  the  depression  of  the  freez- 
ing point  of  the  sap  with  cold  resistance. 

Chandler20  directed  much  attention  to  the  relation  of  osmotic 
concentration  to  hardiness,  although  he  admitted  that  the  force  of 
imbibition  may  be  the  more  important  factor  in  the  water-retaining 
power  of  plant  tissue.  He  found  in  most  cases  that  the  hardier  plants 
had  the  more  concentrated  sap.  To  explain  the  relation  of  a slight 
difference  in  freezing  point  depression  to  a considerable  difference  in 
hardiness,  Chandler  reasoned  that,  since  in  a solution  containing 
one  gram  molecule  the  freezing  point  is  -1.86° C.,  and  in  a M/2 
solution,  -0.93°C.,  in  the  latter  solution  at  a temperature  of  -0.93°C. 
all  the  water  would  be  unfrozen,  at  -1.86 °C.  one  half  would  be  un- 
frozen, and  at  -3.72 °C.  one-fourth  would  be  unfrozen,  and  so  on. 
If  this  held  true  for  the  water  contained  in  a plant,  the  sap  of  which 
is  equivalent  to  about  one-half  gram  molecular  concentration,  we 
would  then  expect  75  percent  of  the  water  to  be  frozen  at  -3.72 °C. 
However,  Chandler’s  conjecture  on  this  point  does  not  apply  in  all 
cases  since  McCool  and  Millar  found  in  their  dilatometer  experiments 
that  nearly  as  much  water  is  frozen  at  -4°C.  in  wheat  plants  having 
a freezing  point  depression  of  1.107 °C.  as  in  corn  plants  having  a 
depression  of  only  0.578 °C. 

Ohlweiler92  in  studying  the  effect  of  a late  spring  frost  on  vege- 
tation at  St.  Louis,  found  that  plants  which  showed  the  greater 
osmotic  concentration  of  the  sap  were  generally  injured  the  least, 
although  there  were  some  exceptions.  He  found,  for  example,  that  in 
twelve  species  of  Magnolia,  the  order  of  hardiness  paralled  the  order 
of  sap  concentration  fairly  well.  Harris  and  Popenoe40  found  that  on 
the  average,  the  hardier  species  of  avocado  had  slightly  the  greater 
sap  concentration.  Lewis  and  Tuttle59  working  on  evergreen  leaves 
in  Canada,  found  that  in  Picea  Canadensis , the  freezing  point  low- 
ering varied  only  slightly  from  October  to  April,  the  maximum 
lowering  being  in  March.  In  the  bark  of  Populns  and  the  leaves  of 
Linnaea  and  Pyrola , the  maximum  depression  of  the  freezing  point 
was  also  found  to  be  in  March,  after  the  coldest  weather  was  over. 
The  freezing  point  depression  was  found  to  parallel  the  accumulation 
of  sugars  during  the  winter  months,  the  maximum  sugar  content 
being  found  April  2nd.,  just  before  spring  growth  started.  They 
found  little  correlation  between  cold  resistance  and  sap  concentra- 
tion, as  measured  by  the  depression  of  the  freezing  point.  Pantanel- 
li95  likewise,  was  unable  to  establish  a relation  between  osmotic  con- 
centration of  the  cell  sap  and  resistance  to  cold. 


The  Hardening  Process  in  Vegetable  Plants. 


17 


Salmon  and  Fleming109  found  no  relationship  between  sap 
concentration  and  winter  hardiness  in  several  common  cereal  crops 
in  Kansas.  Thus  on  November  27th.,  hardy  Kharkov  wheat  gave 
a freezing  point  depression  of  1.230° C.  and  tender  Culberson  oats 
1.199°C.  On  December  17th.,  the  freezing  point  depression  of  the 
wheat  was  0.935°C.  and  of  the  oats  1.260°C.  They  explain  these 
results  by  the  supposition  that  oats  are  less  able  to  secure  sufficient 
water  from  the  soil  to  supply  that  lost  by  transpiration,  the  ground 
being  frozen  at  the  time  of  the  second  determination.  This  resulted 
in  water-depletion  in  the  oat  plants,  giving  a higher  cryoscopic  value 
to  their  sap. 

Wiegand131  thought  osmotic  concentration  of  plant  sap  to  be  of 
importance  in  relation  to  ice-formation  at  the  inception  of  freezing 
only. 

Imbibition  and  icater-retaining  power. — The  term  “imbibition” 
will  be  used  in  this  paper  in  the  general  sense,  as  applying  to  the 
absorption  of  water  by  colloidal  materials  and  the  holding  of  water 
by  finely  divided  solids  by  means  of  surface  phenomena,  such  as 
adsorption,  adhesion  or  molecular  capillarity. 

De  Candolle75  (quoted  by  Lindley  in  1855)  formulated  the  fol- 
lowing laws  of  temperature  in  relation  to  plants : 

“1.  The  power  of  the  plant  to  resist  low  temperature  is  in 
inverse  ratio  of  the  water  content. 

“2.  Hardiness  is  in  direct  proportion  to  the  viscidity  of  the 
plant’s  fluids. 

“3.  Hardiness  is  in  inverse  ratio  to  the  rapidity  with  which  the 
fluids  circulate. 

“4.  Tenderness  is  greater  in  proportion  to  the  size  of  the  cells.” 

Considering  that  De  Candolle  had  few  or  no  experimental  data 
from  which  to  draw  conclusions,  and  that  he  wrote  many  years  be- 
fore the  classical  researches  of  Miiller-Thurgau,  his  views  on  the 
resistance  of  plants  to  low  temperature  are  remarkably  near  present 
conceptions. 

Wiegand131  considered  that  the  force  of  imbibition  was  to  a 
large  extent  the  cause  of  the  water-retaining  power  of  plant  cells. 
According  to  Pfeffer140  this  force  increases  with  decreasing  moisture 
content.  Although  Wiegand  made  no  quantitative  measurements, 
his  theories  were  the  result  of  keen  observation  and  sound  reason- 
ing and  are  of  very  great  importance  to  an  understanding  of  the 
differential  killing  of  plants  by  cold.  He  pointed  out  that  the  water 
of  crystallization  in  frozen  plant  tissue  was  practically  pure,  sepa- 


18 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


rating  from  the  other  cell  constituents  upon  freezing.  The  progres- 
sive dehydration  of  the  cell  by  the  withdrawal  of  water  to  form  ice 
crystals,  was  thought  by  Wiegand  to  increase  the  combined  forces 
of  osmosis  and  imbibition  holding  the  remaining  molecules  of  water. 
He  advanced  the  hypothesis  that  the  degree  of  cold  necessary  to  form 
ice  was  proportional  to  the  force  which  held  the  w^ter  in  the  tissues, 
which  force  (osmosis  plus  imbibition)  was  thought  to  depend  largely 
on  the  water  content.  Wiegand  believed  that  in  succulent  tissues  of 
high  water-content,  most  of  the  water  would  be  frozen  out  near  the 
initial  freezing  point  and  a smaller  portion  would  be  frozen  in  less 
succulent  tissues. 

Wiegand129  observed  that  no  apparent  ice  formation  took  place 
in  the  buds  of  Quercus,  Castanea,  Hicorea,  Juglans , and  Fraximus, 
at  -18 °C.  The  buds  of  these  species  were  observed  to  differ  from 
many  others  in  which  ice  formation  took  place  at  a higher  tempera- 
ture by:  (1)  lower  water  content,  (2)  smaller  cells,  (3)  thicker  cell 
walls.  He  considered  that  these  factors  favored  the  retention  of  cell 
moisture  by  a relatively  greater  force  of  imbibition  than  in  buds  lack- 
ing such  characteristics  and  in  which  ice  forms  at  a higher  tempera- 
ture. Wiegand  also  observed  that  the  ice  crystals  in  frozen  beets 
and  potatoes  were  smaller  near  the  periphery  than  in  the  center  of 
these  organs.  The  cells  of  the  peripheral  regions  in  these  roots  being 
smaller  and  poorer  in  water,  were  thought  to  have  a greater  capacity 
for  retaining  water  against  the  formation  of  ice  crystals. 

Recent  work  by  Parker97  strengthens  Wiegand ’s  hypothesis  that 
decreasing  water  content  increases  the  force  of  imbibition.  He  found 
that  finely  divided  materials  in  suspension  held  a considerable  amount 
of  water  as  capillary  surface  films,  and  the  force  with  which  this 
capillary  water  was  held  increased  rapidly  with  decreasing  moisture 
content.  That  moisture  content  has  a marked  influence  on  the  force 
of  imbibition  is  indicated  also  by  the  work  of  Reinke,139  who  found  that 
a pressure  of  sixteen  atmospheres  would  squeeze  water  from  a frond 
of  Laminaria  when  the  moisture  content  was  73  percent,  but  when 
the  moisture  content  was  reduced  to  48  percent,  it  required  a pres- 
sure of  200  atmospheres  to  extract  water. 

If  decreasing  moisture  content  increases  the  force  with  which 
water  is  retained  by  plant  cells,  a direct  connection  is  indicated  be- 
tween such  water-retaining  power  and  cold  resistance,  for  several 
investigators  working  with  a wide  variety  of  plants  have  shown  that 
hardiness  is  usually  associated  with  low  moisture  content.  Thus, 
Lindley61  recognized  the  fact  that  decreasing  the  moisture  content 


The  Hardening  Process  in  Vegetable  Plants. 


19 


tended  to  increase  cold  resistance  and  that  the  removal  of  some 
water  in  the  “ripening  process’’  made  the  plant’s  tissues  better  able 
to  withstand  cold.  Detmer26  stated  that  such  parts  of  plants  as  are 
poor  in  water  withstand  low  temperature  best.  He  found  that  air- 
dry  seeds  of  Triticum  and  Pisurn  germinated  normally  after  exposure 
to  temperature  of  -5°  to  -10°  C.,  while  turgid  seeds  were  killed  under 
the  same  conditions. 

Gorke35  noted  that  the  more  hardy  plants  had  the  greater  per- 
centage of  dry  matter  and  slightly  lower  sap  freezing  point.  Schaff- 
nit110  found  a gradation  in  the  amount  of  dry  matter  in  different 
varieties  of  wheat  in  direct  proportion  to  their  resistance  to  low 
temperature.  He  concluded  that  high  dry-matter  content  was  cor- 
related with  high  frost  resistance.  Rivera103  found  that  all  cultural 
conditions  which  tended  to  increase  the  percentage  of  dry  matter  in 
wheat  decreased  the  tendency  to  lodging  and  increased  hardiness. 
Hedlund44  found  that  under  like  cultural  conditions,  those  varieties 
of  winter  wheat  having  a higher  percentage  of  dry  matter  in  autumn 
are  generally  more  winter-hardy  than  those  having  a low  percentage. 
He  found  also  that  cultural  conditions  that  make  for  high  percentage 
of  dry  matter  favor  winter  hardiness.  Hedlund  attributed  the  high 
dry-matter  content  of  hardy  plants  to  their  large  carbohydrate  con- 
tent. 

Shutt114  found  that  a correlation  existed  between  percentage  of 
dry  matter  and  hardiness  in  apple  twigs.  A set  of  samples  gathered 
on  the  Canadian  Experiment  Farm  in  midwinter  had  moisture  con- 
tents ranging  from  45.1  percent  in  terminal  parts  of  twigs  of  Yellow 
Tiansparent  (hardy)  to  51.59  percent  in  the  same  portion  of  the 
Blenheim  Pippin  (tender).  He  recommended  the  use  of  cultural 
practices  to  regulate  the  moisture  content,  as  indicated  by  the  degree 
of  maturity  in  the  fall.  It  is  now  a pretty  well  recognized  fact  that 
the  ability  of  a variety  of  the  apple  to  survive  in  Northern  sections 
depends  on  its  maturing  thoroughly  before  winter — in  other  words, 
developing  a condition  of  low  moisture  content  and  maximum  water 
retaining  power.  Webber127  and  his  co-workers  observed  after  a very 
severe  freeze  in  the  citrus  regions  of  California  that  trees  and  por- 
tions of  trees  which  were  dormant  or  inactive  were  much  less  injured 
than  those  actively  growing  and  functioning.  Trees  which  had  been 
rather  dry  for  some  time  also  were  more  hardy  than  those  recently 
irrigated  while  trees  suffering  badly  from  drought  were  injured  worst. 

Batchelor  and  Reed5  found  that  winter-injury  of  the  distal  end 
of  the  branches  of  the  Persian  walnut  in  California  could  be  pre- 


20 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


vented  by  bringing  the  trees  to  early  maturity  by  with-holding  water, 
followed  with  heavy  irrigation  during  the  winter. 

Johnson51  found  a marked  seasonal  increase  in  water  content  of 
peach  buds  in  Maryland,  correlated  with  the  increased  tenderness  of 
buds  in  spring.  The  variety  Greensboro  had  a lower  water  content 
than  the  Elberta,  which  is  a tenderer  variety.  West  and  Edlefsen128 
also  working  on  peach  buds,  pointed  out  that  buds  might  escape 
injury  from  cold  by  under-cooling  below  the  freezing  point  without 
ice  formation,  when  the  amount  of  moisture  in  the  buds  was  small. 

Chandler20  and  more  recently  Carrick18  found  that  apple  roots 
which  had  been  allowed  to  absorb  moisture  for  several  hours  were 
injured  by  cold  a little  more  than  normal  roots,  whereas  partial  drying 
increased  their  cold  resistance. 

Beach  and  Allen6  found  that  drying  apple  twigs  before  freezing 
lessened  the  injury  by  cold.  They  also  found  that  the  hardier  va- 
rieties of  apples  have  the  lower  moisture  content  during  the  growing 
season  but  after  prolonged  freezes  in  winter,  these  hardy  sorts  may 
contain  more  moisture  than  tender  varieties.  In  other  words,  the 
hardy  twigs  undergo  a smaller  water  loss  during  freezing. 

Salmon  and  Fleming109  performed  an  interesting  experiment 
with  greenhouse-grown  cereal  plants,  which  demonstrated  that  cold 
resistance  may  be  increased  by  decreasing  the  amount  of  water  in  the 
tissues  by  slight  wilting.  Wheat  plants  were  dug  up,  wilted  for  two 
or  three  hours,  and  exposed  to  freezing  temperatures.  Turgid  plants 
killed  much  worse  than  slightly  wilted  plants  at  a temperature  of 
-2  to  -3°C.  for  20  to  30  minutes. 

Chandler20  compared  the  relative  extent  of  killing  by  cold  in 
turgid  and  wilted  plants.  He  included  in  his  experiments  a large 
number  of  tender  plants  which  are  incapable  of  withstanding  ice 
formation  and  which  cannot  be  expected  to  show  much  response  in 
the  way  of  hardiness  to  any  treatment.  His  experiments  were  made 
in  summer,  hence  the  killing  at  temperatures  only  slightly  below 
freezing.  Though  Chandler  concluded  that  on  the  whole,  wilting 
does  not  increase  cold  resistance,  yet  the  following  table,  taken 
from  his  data,  indicates  that  under  certain  conditions,  wilting  may 
do  so. 

In  the  case  of  lettuce,  it  seems  that  the  wilted  plants  were  killed 
the  worst  by  slight  freezing,  -2°C.  At  the  lower  temperatures,  how- 
ever, the  percentage  killed  increases  very  rapidly  in  the  turgid  plants, 
and  slowly  in  the  wilted  plants,  so  that  the  killing  of  turgid  leaves 
considerably  exceeds  that  of  the  wilted  when  the  temperature  of 


The  Hardening  Process  in  Vegetable  Plants. 


21 


Effect  of  Wilting  on  Killing  by  Cold,  Compiled  from  Chandler,  p.  196. 


Plant 

Condition 

Temperature 

-2°C. 

-3°C. 

-*&C. 

-4.5°C. 

Lettuce 

turgid 

wilted 

12y2%  killed 
47%  killed 

66.6% 

55.5% 

83% 

62% 

Red  Clover 

turgid 

wilted 

17%  killed 
34%  killed 

100% 

66.6% 

Rose  Geranium 

turgid 

wilted 

97%  killed 
60% 

100% 

100% 

Red  Cabbage 

turgid 

wilted 

65% 

44% 

-4.5 °C.  is  reached.  The  same  thing  is  indicated  in  the  case  of  red 
clover.  Chandler  remarks  that  brief  wilting  does  not  increase  the 
total  amount  of  material  in  the  cell  sap  which  might  function  in  hold- 
ing water  in  solution,  yet  it  seems  that  the  hardiness  of  the  plants 
may  be  materially  affected. 

Wiegand131  states  that  the  greater  the  water  content,  the  thicker 
the  film  of  water  on  the  surface  of  an  imbibing  substance,  such  as  the 
plant  cell,  and  the  weaker  the  force  by  which  the  outer  layers  of  this 
film  are  held,  hence  more  easily  withdrawn  to  form  ice.  Parker07 
has  furnished  some  experimental  data,  which  substantiates  Wiegand ’s 
suggestion. 

Kiesselbach  and  Ratcliff52  in  experiments  with  seed  corn,  found 
that  death  from  freezing  was  directly  proportional  to  the  moisture 
content  of  the  kernel  and  to  the  duration  of  exposure  to  cold.  Seed 
corn  maturing  in  the  natural  way  was  found  to  become  cold-resistant 
progressively  as  the  moisture  content  decreased.  The  following  table 
taken  from  their  data,  illustrates  the  relation  between  moisture  con- 
tent and  killing  by  cold  as  measured  by  the  germination  of  the  seed. 

Kiesselbach  and  Ratcliff  found  that  the  temperature  as  which  ice 
formation  commences  in  the  corn  kernel  depends  very  largely  on  the 
moisture  content.  Immature  seed  containing  60  to  80  percent  mois- 
ture, froze  just  below  32 °F.,  whereas  in  air-dry  seed,  containing  18 
percent  moisture,  no  ice  formation  could  be  detected  at  -10°F.  Usual- 
ly where  ice  formation  took  place  in  the  seeds  and  they  remained  in 
the  frozen  condition  24  hours,  the  vitality  was  weakened  or  destroyed, 
but  in  some  cases  ice  formation  within  the  seed  was  not  followed  by 


22 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Relative  Germination  of  Seed  Corn  of  Varying  Moisture  Content  After 
Exposure  to  Low  Temperatures.  (After  Kiesselbach  & Ratcliff) 


Temperature 

Percent 

moisture  content 

of  grain 

to  which 

exposed 

Degrees  F. 

10 

15 

20 

25 

30 

35 

40 

45 

50 

60 

to 

to 

to 

to 

to 

to 

to 

to 

to 

to 

15 

20 

25 

30 

35 

40 

45 

50 

55 

65 

32—28 

100 

85 

75 

71 

69 

— 

33 

0 

24—20 

100 

96 

77 

67 

13 

12 

12 

6 

0 

16—12 

100 

88 

34 

12 

0 

0 

0 

0 

0 

8—4 

100 

98 

47 

7 

0 

0 

0 

0 

0 

0 

0—  -5 

97 

63 

0 

0 

0 

0 

0 

0 

0 

0 

death.  They  show  that  air-dry  seed  are  uninjured  by  low  tempera- 
ture, and  that  ice-formation  does  not  take  place  therein. 

The  observations  of  G-orke,  Schaffnit,  Rivera,  Hedlund,  Shutt, 
Webber,  Wiegand,  Beach  and  Allen,  West  and  Edlefsen,  Batchelor 
and  Reed  and  Johnson,  indicate  that  individual  plants,  species  or 
varieties  having  a low  moisture  content  are  usually  hardier  to  cold 
than  those  having  a high  moisture  content.  The  work  of  Chandler, 
Carrick,  Beach  and  Allen,  Salmon  and  Fleming,  and  Kiesselbach  and 
Ratcliff  indicates  that  reducing  the  moisture  content  of  a given 
plant  or  part  of  a plant  increases  its  cold  resistance.  This,  it  seems, 
may  be  partly  accounted  for  by  Wiegand ’s  hypothesis  and  Parker’s 
recent  work,  in  that  the  force  with  which  water  is  held  by  plant  cells 
increases  with  decreasing  water  content.  Removal  of  some  water 
by  drying  before  freezing  should  increase  the  force  with  which  the 
remaining  moisture  is  held.  In  other  words,  if  plant  tissues  become 
more  cold  resistant  upon  slight  drying  out,  such  increase  in  hardiness 
may  be  ascribed  to  the  increased  power  of  imbibition  on  the  part  of 
the  plant’s  cells. 

Relation  of  factors  influencing  water  loss  by  the  plant  as  a 
whole,  to  hardiness. — The  foregoing  discussion  has  shown  the  re- 
lation of  some  factors  to  the  water-retaining  power  of  plant  tissue, 
as  measured  by  the  effects  of  low  temperature.  It  is  indicated  that 
increasing  the  water-retaining  power  of  the  cell,  either  by  increasing 
the  concentration  of  its  sap,  or  by  increasing  its  power  of  imbibiton, 
or  both,  results  in  greater  resistance  to  low  temperature  because  of 
the  increased  force  of  crystallization  necessary  to  withdraw  the  re- 
quired amount  of  water  to  cause  death  or  bring  about  the  changes 
which  cause  death.  If  the  ability  of  the  individual  cell  to  retain  some 
moisture  when  exposed  to  freezing  is  the  significant  point  of  differ- 


The  Hardening  Process  in  Vegetable  Plants. 


23 


ence  between  tender  and  hardy  tissues,  then  the  plant  as  a whole 
may  show  the  same  difference  in  water-retaining  power  and  resistance 
to  water  loss,  but  this  does  not  imply  necessarily  that  hardiness  and 
drought  resistance  go  together.  Salmon108  remarks  that  some  hardy 
grasses  thrive  best  in  damp  localities.  In  drought-resistant  species, 
the  plant  as  a whole  may  be  protected  against  water-loss  by  morpho- 
logical differences  in  structure,  such  as  special  water  storage  tracts, 
few  or  small  stomata,  thick  integument,  bark,  scales,  xerophytic 
characters  in  general;  yet  the  individual  cell  may  possess  little 
water-retaining  power  which  would  prevent  the  excessive  withdrawal 
of  water  upon  freezing. 

While  a low  transpiration  rate  due  to  morphological  modifica- 
tions would  undoubtedly  be  of  great  assistance  to  plants  in  withstand- 
ing injury  from  physiological  drought,  a low  transpiration  rate  also 
may  be  associated  with  high  water  retaining  power  of  the  cells. 

Beach  and  Allen6  observed  a loss  of  four  to  nine  percent  in 
weight  of  apple  twigs  during  a single  week  in  January  with  a mini- 
mum temperature  of  -15°F.  They  found  that  in  general  the  hardiest 
varieties  are  most  resistant  to  the  loss  of  water. 

Strausbaugh117  found  that  coincident  with  the  breaking  of  the 
rest  period  in  semi-hardy  varieties  of  the  plum  in  midwinter,  the 
moisture-retaining  power  of  twigs  and  buds  decreased  rapidty,  while 
in  the  hardy  variety  Assiniboine,  which  remained  dormant  until  early 
spring,  the  water-retaining  power  remained  constant.  This  is  sig- 
nificant, since  increased  tenderness  to  cold,  especially  of  the  flower 
buds,  follows  the  break  of  the  winter  rest. 

Sinz115  concluded  as  a result  of  experiments  at  the  University  of 
Goettingen  that  those  varieties  of  winter  wheat  which  seemed  able 
to  prevent  rapid  transpiration,  were  among  those  most  highly  re- 
sistant to  cold. 

Weaver  and  Morgensen126  in  Nebraska  found  that  in  winter  the 
water  losses  of  coniferous  trees  with  their  needles  intact,  are  relative- 
ly no  greater  than  are  the  losses  from  deciduous  trees  after  leaf-fall. 
This  indicates  great  water-retaining  capacity  in  the  foliage  of  coni- 
fers, most  of  which  are  very  hardy. 

Some  writers  have  likened  hardy  to  desert  plants  because  of  their 
xerophytic  characters,  by  which  water  loss  is  reduced  to  a minimum. 
Thus  Schimper112  states  that  desert  plants  frequently  have  a strong 
resemblance  in  their  structure  and  habit  of  growth  to  those  of  polar 
regions,  as  would  be  expected  if  resistance  to  cold  depended  on  the 
reduction  of  water  loss  to  a minimum.  What  Schimper  probably 


24 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


had  in  mind  was  the  form  of  injury  due  to  physiological  drought, 
where  above-ground  plant  tissues  are  killed  by  desiccation  resulting 
from  their  inability  to  obtain  water  from  a frozen  soil  or  through  a 
frozen  stem. 

Storber118  states  that  “ winter  leaves’  ’ of  herbs  are  quite  xero- 
phytic  in  structure,  enabling  them  to  survive  the  severe  conditions 
to  which  they  are  exposed.  He  points  out  a fact  that  seems  to  have 
been  hitherto  overlooked — that  the  low  water  content  and  high  os- 
motic concentration  in  hardy  plants  may  insure  to  them  more  ready 
absorption  of  soil  water.  This  would  certainly  be  of  great  impor- 
tance to  plants  in  winter,  in  overcoming  physiological  drought,  as 
well  as  increasing  the  resistance  to  the  direct  effects  of  freezing. 
Dachnowski22  observed  xerophytic  developments  in  plants  exposed  to 
physiological  drought  conditions  in  bogs.  Modifications  were  found 
enabling  certain  plants  to  survive  in  bogs  in  spite  of  slow  water  ab- 
sorption due  to  toxicity  of  bog  waters.  The  following  are  the  chief 
modifications  to  which  Dachnowski  ascribes  resistance  to  rapid  wTater 
loss  in  leaves  of  bog  plants. 

1.  Reduction  in  size  of  leaves. 

2.  Thick- walled  epidermis. 

3.  Cuticle,  wax,  and  hairs. 

4.  Mucilaginous  and  resinous  bodies  in  leaves  and  roots. 

Groom36  stated  that  the  function  of  mucilages  and  tannin  in 

buds  is  to  help  hold  the  water  in  the  young  shoots.  Chandler20  found 
that  the  bud  scales  of  the  peach  had  no  influence  on  the  resistance 
of  the  embryonic  tissue  to  low  temperature,  but  that  they  served  as 
protection  against  drying  out  by  repeated  freezing  and  thawing. 
Wiegand130  recognized  that  loss  of  water  from  the  plant  might  take 
place  by  evaporation  from  the  ice  masses  in  frozen  tissues,  and  sug- 
gested that  bark  and  bud  scales  serve  as  protection  against  such  loss. 
As  pointed  out  earlier  in  connection  with  the  rate  of  thawing,  pro- 
tection against  such  loss  of  water  would  be  most  important  in  tis- 
sues exposed  to  repeated  freezing  and  thawing,  as  buds  undoubtedly 
are  in  winter. 

In  a number  of  recent  experiments  on  the  raspberry  in  Ne- 
braska, Emerson*  found  that  by  coating  the  canes  with  paraffin, 
winter-injury  could  be  prevented.  He  observed  that  untreated  canes 
killed  only  to  the  snow-line.  Emerson’s  results  indicate  that  me- 
chanical protection  against  loss  of  water  by  the  plant  as  a whole, 

* Emerson,  R.  A.  Cornell  University,  Ithaca,  New  York,  Personal  correspondence  with 
F.  C.  Bradford. 


The  Hardening  Process  in  Vegetable  Plants. 


25 


may  prevent  the  form  of  winter  injury  due  to  local  physiological 
drought,  wherein  parts  of  plants  exposed  to  repeated  freezing  and 
thawing  and  consequently  to  loss  of  water  which  cannot  Le  replaced 
because  of  frozen  stem  or  frozen  or  dry  soil,  are  eventually  kb  led  by 
the  progressive  desiccation  of  the  tops.  This  type  of  cold  injury 
is  distinct  from  the  direct  effects  of  low  temperature,  }et  some  of 
the  factors  which  increase  the  water-retaining  power  of  the  tissues 
in  the  latter  case  may  also  be  of  importance  in  enabling  the  plant 
to  withstand  the  former. 

Irmscher49  attempted  to  correlate  the  cold  resistance  of  certain 
peat  mosses  with  their  ability  to  withstand  long  drying  out.  He 
found  that  most  species  could  stand  a temperature  as  low  as  -20° C., 
but  they  were  all  killed  at  -30°C.  He  states  that  “no  thoroughgoing 
parallel  was  found  between  cold  resistance  and  ability  to  survive 
long  slow  drying.’ ’ However,  he  found  that  any  particular  species 
could  be  made  more  resistant  to  frost  by  previous  drying  out.  Mosses 
growing  in  a dry  location  were  found  more  hardy  than  the  same 
species  in  moister  places.  Irmscher  attributed  to  a 1 1 regenerative 
cell-complex”  the  means  by  which  these  mosses  were  enabled  to 
survive  both  extreme  cold  and  extreme  drying.  A higher  osmotic 
concentration  and  greater  cold  resistance  was  observed  in  species  of 
moss  growing  at  low  temperature. 

STATEMENT  OF  PROBLEM. 

The  work  of  the  earlier  investigators  shows  that  freezing  to 
death  of  plant  tissue  is  associated  with  water-withdrawal  from  the 
cells — the  actual  death  process  being  due  to  (a)  the  direct  effect  of 
water  subtraction  on  the  protoplast,  or  (b)  precipitation  of  proteins 
because  of  the  increased  acidity,  or  (c)  precipitation  of  proteins 
due  to  increased  salt  concentration,  or  perhaps  to  other  processes 
which  have  not  as  yet  received  attention. 

Regardless  of  the  particular  theory  which  may  account  for  the 
ultimate  killing  of  plant  tissue  by  cold,  the  consideration  that  the 
primary  factor  is  water-withdrawal  logically  suggests  the  following 
questions.  In  general,  would  not  cold  resistance  be  proportional  to 
the  water-retaining  capacity  of  the  plant  cells?  Since  the  force  of 
imbibition  increases  with  decreasing  moisture  content  and  since  also 
cold  resistance  in  plants  increases  with  decreasing  moisture  content, 
does  not  cold  resistance  depend  largely  on  the  imbibitional  fo'  ce  with 
which  the  cells  retain  moisture?  Do  hardy  plant  cells  actually  re- 
tain more  moisture  when  exposed  to  freezing  than  cells  of  tender 


26 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


plants?  Do  tender  plants  exposed  to  hardening  treatments  acquire 
an  increased  cell-water-retaining  power,  and  if  so,  is  this  the  main 
factor  concerned  in  their  increased  cold  resistance?  Also,  how 
is  this  increased  water-retaining  power  acquired  and  what  changes 
in  the  living  plant  are  concerned  therein?  In  order  to  answer  these 
questions,  the  following  experimental  work  has  been  undertaken. 

EXPERIMENTAL  WORK. 

Materials  used. — Most  of  the  experiments  were  performed  with 
the  cabbage,  as  a representative  of  a type  of  plant  which  is  capable 
of  being  hardened  so  as  to  withstand  considerable  ice  formation 
within  the  leaves.  Leaf  lettuce,  head  lettuce,  kale,  cauliflower  and 
celery  were  used  to  some  extent.  These  also  are  plants  capable  of 
being  hardened  so  that  they  can  be  frozen  stiff  without  injury. 

The  tomato  was  used  as  the  principal  representative  of  a type  of 
plant  which  cannot  be  hardened  so  as  to  withstand  ice-formation, 
but  which  is  capable  of  hardening  to  the  extent  that  the  freezing 
point  is  lowered  slightly.  Other  plants  used  of  this  type  were  pep- 
pers, eggplant  and  sweet  potatoes. 

In  each  series  of  experiments  plants  of  the  same  variety  and 
age  were  used. 

Methods  of  hardening. — Series  E. — The  plants  were  kept  in  a 
warm  greenhouse  until  nearly  large  enough  for  transplanting  to 
the  garden.  The  plants  to  be  hardened  were  then  removed  to  an 
open  coldframe  where  they  were  exposed  to  temperatures  near 
freezing  during  the  night  and  to  full  sunlight  during  the  day.  This 
method  of  hardening  was  followed  both  in  early  spring  and  in  late 
fall.  Samples  were  gathered  for  analysis  usually  at  intervals  of 
5,  10  and  20  days  after  the  beginning  of  the  hardening  treatment, 
as  well  as  from  some  of  the  original  lot  of  plants  which  had  been 
kept  in  the  greenhouse  under  favorable  growing  conditions.  The 
soil  moisture  supply  was  kept  as  nearly  as  possible  the  same  for  the 
plants  in  the  greenhouse  and  those  being  hardened  in  the  frames, 
so  that  temperature  would  be  the  principal  limiting  factor  in  their 
development. 

Series  A. — The  soil  moisture  for  plants  grown  in  a warm  green- 
house was  varied.  As  soon  as  the  seedlings  were  well  established 
after  transplanting  from  the  seed  flat,  a number  of  potted  plants  of 
uniform  size  were  selected  and  divided  into  lots  which  were  given 


The  Hardening  Process  in  Vegetable  Plants. 


27 


different  treatment  only  in  so  far  as  water  supply  was  concerned. 
One  lot,  Al,  was  given  liberal  moisture — these  plants  were  kept  in 
rapidly  growing  condition  and  were  always  the  tenderest  plants  in 
the  experiments.  Another  lot,  A2,  was  given  moderate  moisture,  so 
that  the  plants  grew  at  a moderate  rate.  Another  lot,  A3,  was  given 
just  enough  water  to  keep  the  plants  growing  slowly.  They  frequent- 
ly wilted  somewhat  in  the  middle  of  warm,  bright  days.  This  lot 
usually  showed  nearly  the  same  degree  of  hardiness  as  those  plants 
that  had  received  the  maximum  degree  of  hardening  in  the  cold- 
frame.  A fourth  lot,  A4,  was  included  in  some  of  the  experiments, 
these  plants  being  watered  liberally  at  first,  then  water  was  partially 
withheld  for  a week  or  ten  days  before  samples  were  taken. 

Series  B and  C. — Plants  were  grown  under  uniform  conditions 
in  the  greenhouse,  in  soils  of  different  composition  made  up  by  mix- 
ing different  proportions  of  sand  and  compost.  Few  data  are  re- 
ported on  this  series  because  it  was  found  difficult  to  maintain  uni- 
form moisture  conditions  in  soils  of  such  diverse  texture.  Also 
other  factors,  such  as  degree  of  root  binding,  were  likely  to  be- 
come limiting  before  excess  or  deficiency  of  nutrients  could  exert 
much  effect.  However,  it  was  definitely  shown  that  growing  plants 
in  poor  soils  would  increase  their  cold  resistance,  other  conditions 
being  the  same.  Such  plants  were  smaller  and  grew  more  slowly 
than  the  more  tender  plants  in  the  better  soils.  This  series  of  ex- 
periments might  have  been  more  successful  if  the  plants  had  been 
grown  in  a uniform  soil-medium  to  which  varying  quantities  of 
nutrient  solution  were  added. 

Series  H. — The  treatment  consisted  of  severely  pruning  the 
roots  by  running  a knife  close  to  the  stem  on  one  or  both  sides  of 
the  plant.  This  treatment  checked  the  growth  of  the  plants  quite 
materially  for  a short  time  and  increased  the  cold-resistance  some- 
what. 

Series  F. — A quite  effective  method  of  hardening  was  watering 
with  M/10  solutions  of  various  salts.  The  plants  were  grown  under 
uniform  conditions  in  a warm  greenhouse  and  the  test  lots  were 
watered  with  the  various  salt  solutions  whenever  the  soil  became 
rather  dry  or  whenever  the  plants  wilted  badly.  In  some  cases,  as 
under  high  transpiration  conditions,  the  wilting  point  was  reached 
while  the  moisture  content  of  the  soil  was  high.  It  is  not  altogether 
clear  whether  the  hardiness  resulting  from  these  salt  applications 
was  due  to  their  specific  action,  to  a condition  of  mild  physiological 


28 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


drought,  or  to  the  toxicity  of  such  concentrated  solutions  to  the  roots. 
This  will  be  discussed  in  more  detail  later. 

EFFECT  OF  HARDENING  TREATMENTS  ON  PLANTS. 

External  appearance.— Cab bage. — Tender  (wet-grown)  green- 
house plants  were  usually  about  twice  as  large  as  those  hardened  by 
withholding  of  water,  as  shown  by  the  relative  green  weights  of  A1 
and  A3  in  Table  2.  Plants  hardened  by  withholding  moisture  were 
usually  darker  green,  covered  with  heavy  waxy  bloom,  with  slight 
pink  tints  in  the  stem  and  petioles,  but  not  as  heavily  pigmented 
as  the  coldframe  hardened  plants.  The  leaves  were  tough  and 
leathery,  in  contrast  to  the  brittle,  crisp  texture  of  tender  plants. 

Cabbage  plants  hardened  by  exposure  in  coldframe  were  smaller 
and  stockier  than  unhardened  greenhouse  plants  and  nearly  always 
showed  more  or  less  pink  pigment  (probably  anthocyanin)  in  the 
stems,  petioles  and  leaf  veins.  Coldframe  hardened  plants  were  tough 
and  leathery  in  texture.  In  most  of  the  experiments  the  maximum 
degree  of  hardening  by  this  method  enabled  cabbage  to  withstand  a 
temperature  of  -5°C.  to  -6°C.  for  at  least  one  hour,  whereas  non- 
hardened  plants  would  be  killed  between  -3°C.  and  -4°C.  In  a few 
experiments  hardened  cabbage  withstood  temperatures  as  low  as 
-8°C.  to  -10°C.  over  night. 

The  development  of  pink  color,  especially  in  the  stems  and  peti- 
oles, was  conspicuous  in  all  hardened  plants.  According  to  Knud- 
son55  the  “work  of  Ewart,  Overton,  Wheldale  and  others  indicates 
a close  relationship  between  the  sugar  content  of  the  plant  and  pig- 
ment production.  ” Throughout  Knudson’s  experiments  on  the  ef- 
fect of  carbohydrates  on  green  plants,  a tendency  to  anthocyanin  pro- 
duction was  observed,  plants  fed  on  glucose  and  maltose  (M/20 
solutions)  showing  heavy  coloration,  which  disappeared  within  a 
week  when  they  were  placed  in  diffuse  light.  These  results  are  of 
special  interest  in  connection  with  the  large  sugar  content  found  in 
hardened  plants,  discussed  later. 

Nicholas91  was  of  the  opinion  that  “the  production  (in  leaves) 
of  anthocyanin  is  correlated  with  the  formation  of  organic  acids.  The 
connection  known  to  exist  between  oxidation  and  pigmentation  in- 
heres in  the  production  of  these  acids,  accompanied  by  the  formation 
of  the  red  pigment/’ 

The  conspicuous  development  of  the  waxy  bloom  on  cabbage 
plants  has  been  considered  by  Harvey42  of  some  importance  in  rela- 
tion to  cold  resistance  in  that  it  may  permit  the  undercooling  of  the 


The  Hardening  Process  in  Vegetable  Plants. 


29 


leaf  several  degrees  below  the  freezing  point.  He  suggests  that  it 
prevents  the  ‘ ‘ inoculation  ’ ’ of  the  moisture  in  the  leaf  by  droplets  of 
water  freezing  on  the  surface. 

Cabbage  plants  hardened  by  other  methods  showed  much  the 
same  changes  as  did  those  in  the  series  mentioned  above.  In  all 
cases  hardiness  was  in  direct  proportion  to  the  external  changes 
noted.  Wherever  the  growth  of  the  plant  was  materially  checked, 
even  for  a few  days,  hardiness  was  increased  in  proportion  to  the 
checking. 

Cauliflower  and  kale  showed  about  the  same  changes  on  harden- 
ing as  cabbage. 

Leaf  lettuce.  Both  small  potted  plants  and  large  plants  ap- 
proaching maturity  in  the  greenhouse  and  coldframe  were  used. 
The  leaves  become  tougher,  thicker  and  of  more  leathery  texture 
upon  hardening.  Pigmentation  was  not  conspicuous.  When  hard- 
ened by  drying,  the  crinkling  of  the  leaves  was  more  pronounced 
and  the  color  deeper  green. 

Tomato.  Leaves  of  hardened  plants  became  very  dark  green 
with  much  pigmentation  on  the  under  side,  were  much  smaller, 
tended  to  curl  on  the  midrib;  the  stems  and  petioles  became  very 
heavily  pigmented,  tough  and  woody  in  texture.  Hardening  tomato 
plants  in  the  greenhouse  by  any  of  the  methods  of  checking  growth 
had  about  the  same  effect  on  external  appearance.  The  same  was 
true  of  the  coldframe-hardened  plants,  except  that  when  hardening 
was  long-continued  at  low  temperature,  the  lower  leaves  turned 
yellow  and  fell,  until  the  plant  was  nearly  defoliated.  This  is  prob- 
ably similar  to  the  form  of  killing  by  temperatures  above  the  freez- 
ing point  noticed  by  Molisch  and  attributed  by  him  to  the  inhibition 
of  metabolism  by  the  low  temperatures. 

Morphological  difference  in  hardened  plants. — Schaffnit110  was 
unable  to  find  any  structural  differences  in  varieties  of  wheat  vary- 
ing in  degree  of  cold  resistance.  Salmon  and  Fleming109  could  find 
no  difference  in  cell  structure  in  hardy  and  tender  varieties  of  cereals. 
On  the  other  hand,  Briggs10  found  that  the  cells  were  somewhat 
smaller  in  the  pistils  of  hardy  varieties  of  peaches.  Walster124  ob- 
served that  in  barley  grown  at  15°C.,  there  was  greater  lignification 
of  the  xylem  bundles  than  in  plants  grown  at  20° C.  This  would 
make  the  plants  grown  at  the  lower  temperature  stiffer  and  stronger. 

To  determine  whether  the  hardening  process  affects  the  size 
of  the  cells  in  vegetable  plants,  sections  were  made  of  hardened  and 


30 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


not  hardened  cabbage  and  tomato  leaves  and  the  palisade  cells  meas- 
ured. Portions  of  young  leaves  which  had  made  most  of  their  growth 
during  the  process  of  hardening  were  used  in  each  case.  Hence  the 
differences  in  the  cells  here  reported  do  not  represent  an  “ acquired  ” 
condition,  but  differences  in  development  between  plants  under  favor- 
able growing  conditions  and  those  subjected  to  hardening.  Portions 
were  taken  from  corresponding  locations  on  leaves  of  about  the  same 
size,  killed  and  fixed  in  the  usual  way.  Transverse  sections  were 
made  with  the  rotary  microtone,  mounted,  stained  and  measured. 

In  the  tender  tomato  leaf  numerous  large  air  spaces  were  ob- 
served, while  in  hardened  leaves  the  cells  were  more  compactly  ar- 
ranged and  filled  with  starch  grains.  Starch  grains  were  less  plen- 
tiful in  hardened  cabbage  leaves,  but  the  compactness  of  the  cellu- 
lar arrangement  was  marked.  Table  1 gives  the  measurements  in 
two  dimensions  of  the  palisade  cells  and  the  cross-section  area  com- 
puted therefrom.  These  data  are  the  averages  obtained  from  meas- 
urements of  several  different  sections. 


Table  1. — Measurements  of  Leaf  Palisade  Cells  in  Plants  Hardened  and 

not  Hardened. 


Thickness 

Thickness 

Thickness 

Width 

Length 

Area. 

of  whole 

of  pali- 

of paren- 

of 

of 

sq.  m- 

leaf 

sade 

chyma 

cells 

cells 

Cabbage 

Not  hardened  . 
Hardened  by- 

....291m 

134.5m 

106. 3m 

19.1m 

36.3m 

694.0 

drying  in  g.  h. 

... .269 

127.9 

106.3 

19.4 

27.8 

538.8 

Not  hardened  . . 

274 

136.2 

102.1 

20.9 

36.9 

772.0 

Hardened  in 
coldframe  

....312 

JL68.6 

118.0 

19.9 

31.1 

619.5 

Tomato 

Not  hardened  . 

196.6  ! 

76.2 

76.2 

21.0 

57.5 

1201.5 

Hardened  . . . . 

....133.7  i 

55.6 

68.0  1 

14.1 

44.8 

630.1 

Judging  from  the  data  presented  in  Table  1,  hardened  plants 
are  characterized  by  somewhat  smaller  and  more  compact  palisade 
cells  than  are  non-hardened  leaves  of  the  same  sort.  In  tomato,  leaves 
from  plants  given  hardening  treatments  are  considerably  thinner 
than  tender  leaves,  however,  cabbage  leaves  hardened  in  coldframe 
gained  in  thickness. 

Effect  of  hardening  treatments  on  rate  of  growth. — The  growth 
of  plants  subjected  to  any  of  the  hardening  treatments  was  checked 
in  proportion  to  the  intensity  of  the  treatment.  Data  are  presented 
in  Table  2,  on  samples  gathered  from  lots  of  the  same  age,  grown 


The  Hardening  Process  in  Vegetable  Plants. 


31 


under  otherwise  uniform  conditions,  except  for  the  various  harden- 
ing treatments.  With  the  average  green  weight  of  the  plants  as  the 
criterion,  it  is  evident  that  hardened  plants  are  much  smaller,  indi- 
cating the  extent  to  which  their  growth  had  been  checked.  Thus, 
in  the  series  gathered  March  12,  1920,  wet-grown  greenhouse  cabbage 
plants  (tender)  averaged  23.1  grams,  plants  hardened  by  partial 
withholding  of  moisture  averaged  6.8  grams  and  plants  hardened  in 
coldframes  three  weeks  averaged  7.67  grams.  The  differences  in  dry 
weight  are  not  so  great,  since  the  smaller  and  hardier  plants  pos- 
sessed a larger  percentage  of  dry  matter. 

In  the  tomato,  the  rate  of  growth  could  be  roughly  measured  by 
the  increase  in  height  from  week  to  week.  Accordingly  a number 
in  each  of  the  lots  subjected  to  the  various  treatments  were  measured 
each  week.  The  retardation  of  growth,  when  any  of  the  hardening 
treatments  became  operative,  is  shown  in  figure  1. 


Effect  of  hardening  treatments  on  percentage  of  dry  matter. — 

The  data  given  under  percent  of  dry  matter  in  Table  2 indicate  con- 
siderable increase  of  dry  matter  in  all  of  the  experimental  lots  of 
plants  exposed  to  hardening  treatments.  Conversely,  the  water  con- 
tent decreased  in  hardened  plants,  roughly  in  proportion  to  the  ex- 
tent to  which  their  growth  was  checked  by  the  hardening  treatment. 


32 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


The  possible  significance  of  decreased  water  content  in  relation  to 
the  water-retaining  power  of  the  cell  when  exposed  to  water-loss  by 
freezing  has  already  been  indicated  by  Wiegand  and  more  recently 
by  the  work  of  Chandler,  Salmon  and  Fleming,  Carrick  and  indi- 
rectly by  Parker.  It  may  be  repeated  here  that  decreased  water  con- 
tent would  be  associated  with  increased  force  of  imbibition,  and 
with  increased  concentration  of  the  cell  sap,  which  forces  tend  to 
retain  water  in  the  cell  during  freezing.  It  is  realized  that  the  total 
loss  in  weight  upon  drying  of  leafy  tissue  does  not  truly  represent 
the  actual  water  content  of  the  plant,  but  the  difference  is  probably 
so  small  that  this  loss  is  taken  as  the  moisture  content  throughout 
these  experiments. 

Effect  of  hardening  treatment  on  depression  of  freezing  point. — 

In  several  experiments,  the  freezing  point  depression  of  the  expressed 
sap  of  leaves  of  hardened  and  non-hardened  plants  was  determined 
with  the  usual  Beckman’s  apparatus.  Potted  plants  from  each  ex- 
perimental lot  were  brought  into  the  laboratory  to  insure  having 
fresh  tissue  for  each  determination.  All  of  the  leaves  were  taken 
from  two  or  three  plants  and  ground.  The  sap  was  then  squeezed 
from  the  macerated  pulp  and  duplicate  or  triplicate  freezing  point 
determinations  were  made  at  once.  The  data  given  in  the  column 
“Depression  of  the  Freezing  Point”  in  Table  2 show  that  the  de- 
pression was  somewhat  greater  in  the  hardened  plants,  indicating 
greater  osmotic  concentration  of  their  sap.  Similar  data  have  been 
obtained  by  Chandler  and  by  Harvey,  working  on  the  same  sort 
of  material,  hence  it  was  not  deemed  worth  while  to  make  a larger 
number  of  these  determinations.  The  differences  found  here  in 
freezing  point  depression  are  somewhat  greater  than  those  obtained 
by  Chandler20  and  much  greater  than  those  reported  by  Harvey42 
for  hardened  and  not  hardened  cabbage.  This  is  due  perhaps  to  the 
extremes  in  the  treatments  used  in  these  experiments.  Heavy  water- 
ing made  Series  A1  somewhat  more  tender  than  ordinary  non-hard- 
ened plants  and  Series  A3  attained  maximum  hardiness  through  the 
application  of  the  minimum  amount  of  water  to  keep  the  plants  from 
wilting. 

It  may  be  pointed  out  that  the  increased  sap  concentration  in 
hardened  plants  is  due  probably  to  the  combined  effect  of  the  follow- 
ing factors:  (1)  Decreased  total  moisture  content.  (See  Table  2). 
(2)  Increase  in  the  amount  of  sap  solutes.  Numerous  investigators 
have  found  an  increase  of  soluble  sugars  in  plants  exposed  to  low 


The  Hardening  Process  in  Vegetable  Plants.  33 


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greenhouse 


34  Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


The  Hardening  Process  in  Vegetable  Plants.  35 


Table  2. — (Continued) . 

Date  I Average  I Average  j % of  I Depression 


36  Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


The  Hardening  Process  in  Vegetable  Plants. 


37 


temperature  and,  as  shown  later,  there  is  an  increased  sugar  con- 
tent in  hardened  vegetable  plants.  (3)  Increased  amount  of  water 
held  in  the  absorbed  state  by  the  cell  colloids.  Since  this  absorbed 
water  is  probably  nearly  pure,  the  sap  solutes  of  hardened  plants 
are  held  in  solution  by  a smaller  volume  of  water — hence  the  greater 
concentration. 

Chandler21  found  that  a large  part  of  the  depression  of  the 
freezing  point  in  plant  sap  was  due  neither  to  sugars  nor  to  electro- 
lytes. Recent  work  by  Parker97  showed  that  finely  divided  material 
in  suspension  exerted  considerable  influence  on  the  freezing  point 
and  that  this  depression  increases  rapidly  with  decreasing  moisture 
content.  Though  Parker ’s  work  was  done  with  soils  and  dried,  finely 
ground  inorganic  colloids,  it  may  be  supposed  that  the  organic  col- 
loidal particles  of  plant  protoplasm  have  the  same  property  of  de- 
pressing the  freezing  point.  Parker  attributes  the  lowering  of  the 
freezing  point  by  finely  divided  material  to  the  force  of  “adhesion” 
by  which  films  of  the  liquid  are  held  on  the  surface  of  the  solid  ma- 
terial. The  freezing  point  depression  caused  by  the  finely  divided 
material  decreases  almost  to  zero  in  presence  of  high  moisture  con- 
tent. He  explains  this  by  the  suggestion  that  as  the  amount  of 
liquid  increases  some  of  it  becomes  so  far  distant  from  the  solid  par- 
ticles and  is  so  weakly  held  that  no  depression  of  the  freezing  point 
occurs.  This  is  very  nearly  the  same  as  Wiegand’s  theory  as  to  the 
holding  of  water  in  plant  cells  by  “molecular  capillarity.”  It  may 
be  that  the  increase  in  colloidally -held  water  alone  would  account 
for  much  of  the  depression  of  the  freezing  point  in  hardened  plants. 
Therefore  it  may  be  said  that  the  apparent  increase  in  osmotic  con- 
centration of  the  sap  in  hardened  plants  is  merely  coincident  with 
the  state  of  being  hardy,  rather  than  a cause  of  it. 

EFFECT  OF  HARDENING  ON  ICE  FORMATION  IN  PLANTS. 

Previous  investigations  on  freezing  of  plant  tissue  have  shown 
that  water  is  drawn  from  the  cells  to  form  ice  in  the  intercellular 
spaces.  Some  plants  are  killed  once  this  occurs.  Others  are  capable 
of  withstanding  some  ice  formation,  but  are  killed  at  lower  tempera- 
tures. Unhardened  cabbage  plants  are  killed  by  freezing  at  -3°C. 
to  -4°C.  The  same  plants  after  being  subjected  to  a hardening 
treatment,  may  withstand  a temperature  of  -5°C.  to  -6°C.  or  even 
a few  degrees  lower. 

Miiller-Thurgau74  has  shown  that  by  no  means  all  of  the  water 
freezes  in  plant  tissue  when  exposed  to  temperatures  well  below 


38 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


the  freezing  point.  His  method  of  measuring  the  amount  of  ice  in 
frozen  plant  tissue  was  based  on  observing  the  cooling  of  water  in 
which  tissue  frozen  to  a definite  temperature  was  placed,  calculating 
that  80  calories  are  required  to  melt  one  gram  of  ice.  In  this  way 
he  was  able  to  show  that  the  amount  of  ice  in  an  apple  increased  as 
the  degree  of  freezing  increased.  The  following  table  is  taken  from 
his  data  on  frozen  apples : 

at  -4.5°C.  percent  total  water  frozen  = 63.8 
at  -7.3  percent  total  water  frozen  = 68.2 

at  -8.0  percent  total  water  frozen  = 72.4 

at  -13.0  percent  total  water  frozen  = 74.4 

at  -14.8  percent  total  water  frozen  = 77.4 

at  -15.2  percent  total  water  frozen  = 79.3 

It  appears  from  these  data  that  the  amount  of  water  frozen  at 
each  successive  fall  in  temperature  decreases  fairly  regularly  until 
-13 °C.  is  reached,  but  below  that  temperature  the  rate  of  ice  forma- 
tion increases  somewhat.  The  latter  temperatures  are  far  below  the 
killing  point  for  apples,  which  may  affect  the  results,  and  Miiller- 
Thurgau’s  technique  may  be  open  to  some  experimental  error. 

Miiller-Thurgau  also  made  some  interesting  determinations  of 
the  time-rate  of  ice  formation.  Kohl-rabi  leaves  exposed  to  a tem- 
perature gradually  declining  from  -5°C.  to  -8.25 °C.  froze  at  the 


Fig.  2. — Relation  of  time  to  rate  of  ice  formation  in  100  grams  kohl-rabi  leaves  (ar- 
ranged from  Mttller-Thurgau’s  data). 


The  Hardening  Process  in  Vegetable  Plants. 


39 


end  of  13  minutes,  when  the  temperature  of  the  leaves  was  only 
-4.3 °C.  and  that  of  the  surrounding  air  was  -7.3 °C.  In  the  first 
half  minute  of  freezing  0.69  grams  of  ice  formed  in  100  grams  of 
leaves.  In  the  next  minute,  2.0  grams,  the  following  two  minutes, 
1.5  grams  per  minute,  and  the  next  minute,  0.8  grams  froze.  There- 
after the  amount  of  ice  formed  per  minute  gradually  decreased  un- 
til, at  the  end  of  one  hour  of  freezing,  41.32  grams  of  ice  had  formed 
in  100  grams  of  leaves,  probably  a little  over  50  per  cent  of  the 
total  water  content.  This  experiment  is  illuminating  as  to  the  re- 
lation of  the  time  factor  to  freezing  of  plants.  It  has  been  observed 
that  in  plants,  such  as  kohl-rabi,  which  can  withstand  some  ice  for- 
mation, injury  at  a temperature  near  the  death  point  is  proportional 
to  the  duration  of  exposure.  This  fact  has  been  observed  frequently 
in  experiments  here  and  Harvey42  presents  in  his  paper  an  excellent 
series  of  photographs  illustrating  the  same  thing.  Figure  2 plotted 
from  Miiller-Thurgau ’s  data,  illustrates  graphically  the  rate  of  ice 
formation.  We  may  regard  the  progressive  increase  in  total  amount 
of  ice  and  the  decrease  in  amount  frozen  per  minute  as  being  due  to 
the  balanced  action  of  the  force  of  crystallization  and  the  water-re- 
taining power  of  the  cells,  the  rate  of  ice-formation  approaching  zero 
as  a limit. 

Foote  and  Saxton28  used  the  dilatometer  in  experiments  on  the 
freezing  of  inorganic  colloids  and  were  able  to  show  that  in  such 
materials  water  existed  in  three  forms,  viz.,  free  water,  capillary 
absorbed  water  and  water  of  chemical  combination.  Recently, 
Bouyoucos10  and  McCool  and  Millar  have  made  use  of  the  dilatometer 
to  measure  the  amount  of  water  freezing  in  soils,  in  plants  and  in 
seeds.  McCool  and  Millar  in  their  latest  report,  state  that  the 
amount  of  water  freezing  in  plants  at  -1.5 °C.  decreased  as  the  con- 
centration of  the  sap  increased  (as  measured  by  the  freezing  point 
method).  At  -4°C.  the  amount  of  water  freezing  was  considerably 
more  than  at  -1.5°C.  and  its  correlation  with  the  freezing  point  of 
the  sap  almost  disappeared.  The  following  figures  rearranged  from 
their  report  illustrate  this : 


Crop-plant 

Date 

Freezing  point 
depression 

Amount  of  water  freezin; 
in  5 grams  of  leaves. 
At  -1.5°C.  At  -4°C. 

Wheat 

Nov.  14 

1.107°C. 

0.40cc. 

2.65cc. 

Rye 

May  17 

1.030 

.86 

2.40 

Rye 

Nov.  24 

.928 

.90 

2.50 

Sweet  clover 

Nov.  24 

.906 

1.22 

2.82 

Red  clover 

May  15 

.780 

1.70 

2.70 

Corn 

June  10 

.578 

2.10 

2.90 

40 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Unfortunately  they  did  not  express  their  results  as  percentages 
of  the  total  moisture  content  in  each  of  the  different  plant  tissues, 
hence  it  is  impossible  to  draw  very  definite  conclusions.  Also  noth- 
ing is  stated  as  to  the  source  of  the  material,  whether  greenhouse  or 
field  grown.  One  interesting  point  to  be  noted  here  is  that  wheat 
and  rye,  which  one  would  expect  to  be  more  cold-resistant  than  corn 
or  clover,  show  a smaller  amount  of  water  frozen  at  -1.5 °C.  and 
somewhat  less  at  -4°C.  It  is  rather  surprising  that  such  decided 
variation  in  sap  density  made  so  little  difference  in  the  amount  of 
water  freezing  at  -4°C. 

In  another  series  of  experiments  with  corn  and  barley  McCool 
and  Millar80  showed  that  varying  the  soil  moisture  content  affected 
the  water  relations  in  the  plants.  Freezing  point  depression  and 
the  percentage  of  moisture  in  the  tops  decreased  slightly  in  plants 
grown  with  15.53  percent  soil  moisture,  as  compared  to  those  grown 
with  23.29  percent  soil  moisture.  The  amount  of  water  freezing  at 
-2.5°C.  was  decreased  somewhat,  and  the  amount  freezing  at  -4.5°C. 
was  decreased  considerably  in  plants  grown  on  the  soils  of  lower 
moisture  content. 

Hibbard  and  Harrington45  found  that  the  freezing  point  of  the 
sap  of  roots  and  tops  of  corn  plants  fell  regularly  as  the  moisture 
content  of  the  soil  in  which  the  plants  were  grown  was  decreased. 
The  following  data  from  their  work  illustrate  the  relation  of  soil 
moisture  to  freezing  point  depression  of  sap. 

Percent  moisture  in  soil  Freezing  point  depression 


of  tops 

of  roots 

31 

1.835°C. 

.492°C. 

23 

1.920 

.600 

16 

2.027 

.647 

13 

2.120 

.942 

11 

2.204 

.995 

McCool  and  Millar80  studied  the  effects  of  varying  the  concen- 
tration of  the  soil  solution,  when  the  moisture  content  was  kept  con- 
stant. They  found  a progressive  increase  in  the  freezing  point  de- 
pression of  the  tops  and  roots  of  plants  grown  in  the  greater  con- 
centrations, but  the  amounts  of  freezable  water  showed  little  varia- 
tion. Earlier  experiments  by  the  same  authors78  showed  that  the 
freezing  point  depression  of  both  tops  and  roots  varied  in  the  same 
direction  as  the  concentration  of  the  soil  solution  in  which  the  plants 
were  grown  but  not  in  proportion  to  it.  They  also  studied  the 
effect  of  varying  the  soil  moisture  content,  keeping  the  concentration 
of  the  soil  solution  constant.  Unfortunately,  in  their  experiment 


The  Hardening  Process  in  Vegetable  Plants. 


41 


the  plants  on  the  high  moisture  soils  took  up  a larger  quantity  of 
the  nutrient  salts  so  that  the  concentration  of  the  soil  solution  soon 
became  less  than  in  the  low  moisture  soils.  Therefore,  it  remains 
undetermined  whether  the  effects  of  wet  and  of  dry  soils  on  freezing 
point  depression  and  amount  of  freezable  water  in  plants  grown 
thereon  are  due  to  the  variation  in  the  water  supply,  or  to  variation 
in  concentration  of  the  soil  which  is  involved,  or  to  both. 

Method  of  measuring  amount  of  water  freezing  in  plant  tissues. 

Though  Miiller-Thurgau  was  able  to  obtain  considerable  data  on  this 
subject  by  measuring  the  latent  heat  of  ice  in  frozen  tissues,  his 
method  is  laborious  and  perhaps  open  to  some  experimental  error. 
The  dilatometer  method,  as  described  by  Bouyoucos11,  presents  great 
advantages  in  its  directness,  simplicity,  and  accuracy  for  measure- 
ments at  different  degrees  of  freezing.  The  use  of  the  dilatometer  is 
based  on  the  fact  that  one  gram  of  water  increases  approximately 
one  tenth  of  its  volume  upon  freezing.  It  has  been  used  in  this 
work  essentially  as  described  by  Bouyoucos,  and  by  McCool  and 
Millar. 

A definite  weight  (4-6  grams)  of  fresh  leaves,  is  placed  in  the 
bowl  of  the  dilatometer,  which  is  then  filled  with  petroleum  ether 
(boiling  point  63°C.).  The  dilatometer  is  then  stoppered  with  a 
rubber  cork  through  which  a thermometer  is  placed,  so  that  the  bulb 
is  in  contact  with  the  leaf  tissue  and  the  scale  convenient  for  reading. 
It  was  found  at  the  beginning  of  this  work  that  quicker  and  more 
accurate  results  can  be  obtained  with  plant  tissues,  by  placing  the 
loaded  dilatometer  in  crushed  ice  for  15  to  20  minutes,  to  lower  the 
temperature  of  the  whole  mass  slowly  and  evenly  to  the  neighborhood 
of  0°C.  After  this  preliminary  cooling,  the  dilatometer  is  plunged 
into  an  ice  and  salt  bath,  mixed  in  such  proportions  that  the  tem- 
perature is  slightly  below  that  which  is  desired  in  the  dilatometer. 
Usually  the  temperature  of  the  plant  tissue  in  the  dilatometer  lags 
slightly  above  that  of  the  freezing  mixture.  The  dilatometer  must 
be  kept  perfectly  still  after  it  is  placed  in  the  bath  in  order  to  secure 
uniform  under-cooling  of  the  plant  tissue,  but  the  freezing  mixture 
can  be  gently  stirred  so  as  to  keep  all  parts  of  the  bath  at  uniform 
temperature. 

At  first  the  column  of  petroleum  ether  in  the  graduated  side-arm 
of  the  dilatometer  falls  rapidly  due  to  the  contraction  of  the  contents 
on  cooling.  It  is  usually  necessary  to  add  a little  more  ether 
to  bring  the  column  up  to  the  point  on  the  scale  where  it  can 


42  Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 

be  read  easily.  When  the  thermometer  indicates  that  the  contents 
of  the  dilatometer  have  been  desired  temperature  for  several 
minutes,  the  position  of  the  column  in  the  side-arm  is  read, 
then  solidification  of  the  under  cooled  water  in  the  plant  tissue 
is  caused  by  tapping  the  dilatometer  against  the  sides  of  the 
bath.  As  solidification  of  the  water  takes  place,  the  column  rises 
in  the  side  arm,  slowly  at  first,  then  more  rapidly  and  then  quite 
slowly  for  several  minutes.  It  usually  takes  5 to  10  minutes  to  es- 
tablish equilibrium,  indicating  that  all  the  water  is  frozen  which 
will  freeze  at  that  temperature.  When  the  column  of  ether  in  the 
side  arm  becomes  stationary,  the  reading  is  taken.  The  amount  of 
expansion  as  a result  of  the  freezing  is  the  difference  between  the 
readings  before  and  after  freezing.  In  this  work  the  side  arm  tube 
was  graduated  to  0.01  cc.  and  readings  could  be  made  to  0.005  cc. 
The  expansion  on  freezing  multiplied  by  10  gives  the  number  of  cc. 
of  ice  formed.  In  some  of  the  earlier  experiments  separate  samples 
were  used  for  moisture  and  dry  matter  determinations.  Later  it  was 
found  that  these  determinations  could  be  made  on  the  dilatometer 
sample  after  the  freezing  experiments  had  been  performed.  The 
water  content  of  the  tissues  being  known,  the  percentage  of  the  total 
water  frozen  can  be  calculated  by  dividing  the  number  of  cc.  of  ice 
formed  by  the  total  water  content  of  the  sample. 

Effect  of  temperature  on  amount  of  water  freezing  in  hardened 
and  non-hardened  cabbage  leaves. — Cabbage  leaves  were  used  in 
most  of  the  experiments,  since  these  were  available  in  varying  de- 
grees of  hardiness.  It  was  found  very  difficult  to  secure  rapid 
crystallization  in  hardened  cabbage  leaves  at  a temperature  higher 
than  -3°C.,  so  that  point  was  taken  for  the  minimum  reading.  On 
the  other  hand,  leaves  could  seldom  be  under  cooled  below  -6°C. 
without  ice  formation,  so  that  point  was  taken  as  the  maximum  limit 
of  freezing  in  most  of  the  experiments.  Dilatometer  determinations 
were  made  a number  of  times  with  each  class  of  material  under  ex- 
periment. These  determinations  were  distributed  over  a period  of  sev- 
eral months.  The  samples  were  taken  at  different  times  of  day,  but  in 
each  series  samples  were  taken  at  the  same  time  of  all  the  different 
types  of  material  which  were  being  compared.  Table  3 gives  the 
results  secured  with  leaves  of  non-hardened  greenhouse  cabbage  plants 
thoroughly  hardened  in  coldframe  (9-12  days)  and  of  plants  har- 
dened in  the  greenhouse  by  partial  withholding  of  water  for  two 
weeks  or  more.  Each  figure  represents  an  average  of  several  deter- 
minations, the  individual  determinations  sometimes  varying  as  much 


The  Hardening  Process  in  Vegetable  Plants. 


43 


as  10  percent  due  to  slight  differences  in  the  material  and  to  the 
hour  at  which  the  samples  were  taken. 


Table  3. — Percentage  of  Total  Moisture  in  Cabbage  Leaves  Frozen  at 
Different  Temperatures. 


Previous  treatment 
of  plants 

Percent  of  dry 
matter 

Percent  of  total  moisture  freezing  at 

-3°C. 

-4°C. 

-5°C. 

-6°C. 

Wet-grown  greenhouse 
plants,  (tender)  . . . 

10.29 

49.9 

75.2 

82.1 

84.2 

Dry-grown  greenhouse 
plants,  (hardy)  .... 

12.7 

27.2 

48.9 

62.6 

71.0 

Hardened  in  coldframe 
9-14  days,  (hardy)  . 

14.67 

29.8 

49.6 

: 58.7 

64.°. 

Considerably  more  water  froze  in  the  tender  plants  than  in  the 


hardened  at  each 
temperature.  The 
material  used  for 
the  dry-grown  hardy 
plant  determinations 
perhaps  was  not 
quite  so  uniformly 
hardy  in  its  nature 
as  that  of  the  two 
other  types.  The 
outstanding  feature 
is  the  progressive  in- 
crease in  percentage 
of  total  moisture 
frozen  as  the  tem- 
perature is  lowered. 
This  is  brought  out 
clearly  in  Figure  3 
plotted  from  the 
data  in  Table  3.  The 
increase  becomes  less 
and  less  for  each 
degree  of  tempera- 
ture lowering.  Thus 
in  the  case  of  the 
tender  cabbage,  26.3 
percent  more  water 
freezes  at  -4°C.  than 


Fig.  3. — Relation  of  temperature  to  percent  of  total 
moisture  freezing  in  tender  and  hardened  cabbage  leaves. 


at  -3°C.,  6.9  percent  more  freezes  at  — 5°C.  than 


44 


Missouri  Agr.  Exp.  Sta.  Researcpi  Bulletin  48 


at  -4°C.  and  2.1  percent  more  freezes  at  -6°C.  than  at  -5°C.  In  the 
coldframe  hardened  plants,  19.8  percent  more  water  is  frozen  at  -4°C. 
than  at  -3°C.,  9.1  percent  more  at  -5°C.  than  -4°C.,  and  5.6  percent 
more  at  -6°C.  than  at  -5°C.  Table  3 shows  that  the  percentage  of 
total  moisture  frozen  at  a given  temperature  is  less  in  the  hardened 
plants.  Table  3A,  constructed  from  the  same  data,  on  the  basis  of  100 
grams  fresh  tissue,  shows  that  the  actual  amount  of  water  remaining 
unfrozen  is  greater  also  in  the  hardened  leaves  although  there  is  a 
smaller  total  moisture  content  in  such  tissues  than  in  tender  leaves. 

Table  3a. — Amount  of  Water  in  100  Grams  of  Cabbage  Leaves  Remaining 
Unfrozen  at  Different  Temperatures. 


Treatment  | 

1 

Percent  dry 
matter 

I 

Percent 

moisture 

Grams  water  remaining  unfrozen 
at 

-3°C. 

-4°C. 

-5°C. 

-6°C. 

Wet-grown  greenhouse 

plants,  tender  

; 10.29 

89.71 

34.9 

22.3 

16.1 

14.3 

Dry-grown  greenhouse 

plants,  hardy  

i 12.70 

87.30 

63.5 

44.6 

32.6 

25.3 

Coldframe,  hardened 

for  9-14  days  

; 14.67 

85.33 

59.9 

! 42.9 

35.2 

30.4 

Since  the  percentage  of  the  total  moisture  which  freezes  at  each 
temperature  is  materially  less  in  hardy  than  in  tender  plants, 
and  since  the  actual  amount  of  water  remaining  unfrozen  is  greater 
in  the  hardy  than  in  the  tender  plants,  we  may  safely  assume  that 
the  cells  of  the  hardened  plants  possess  a greater  power  to  retain 
water  when  exposed  to  freezing.  Although  the  amount  of  water 
frozen  increases  with  the  lowering  of  the  temperature,  we  may  as- 
sume that  whatever  the  nature  of  the  water-retaining  force,  it  is 
overcome  in  successively  smaller  increments  by  the  force  of  crystalli- 
zation as  the  temperature  is  lowered.  The  percentage  of  water  re- 
maining unfrozen  in  the  hardened  leaves  is  approximately  a logarith- 
mic function  of  the  temperature. 

The  hardiest  plants  used  in  this  experiment  probably  could  have 
been  killed  by  long  exposure  to  -6°C.  to  -8°C.  However,  it  may  be 
predicted  from  the  rate  of  increase  in  the  amount  of  water  frozen  at 
the  lower  temperatures,  that  if  in  some  way  the  water-retaining 
power  of  the  cells  in  these  plants  was  increased  slightly,  a much 
lower  temperature  could  have  been  sustained.  Maximow  has  shown 
that  sections  of  cabbage  leaves  which  were  injured  at  -5.2°C.  when 


The  Hardening  Process  in  Vegetable  Plants. 


45 


frozen  in  water,  successfully  withstood  a temperature  of  -32° C.  in 
2-mol.  sugar  solution. 

Changes  in  amount  of  freezable  water  during  the  hardening 
process. — Harvey42  stated  that  cabbage  plants  kept  at  3°C.  for  24 
hours  showed  slightly  increased  hardiness  and  at  the  end  of  five  days 
a considerable  degree  of  hardiness  was  developed.  In  the  present 
experiments,  absolutely  controlled  conditions  were  not  available ; 
however,  it  is  generally  considered  that  about  two  weeks’  exposure  of 
greenhouse-grown  cabbage  plants  in  the  open  coldframe  during  March 
will  bring  about  maximum  hardening.  To  study  the  relation  of  the 
amount  of  freezable  water  to  the  hardening  process,  lots  of  tender 
cabbage  plants  were  removed  from  the  greenhouse  to  the  coldframe 
at  intervals  and  dilatometer  determinations  made  on  the  leaves  of 
these  plants  which  represented  progressive  degrees  of  hardening. 
The  results  are  presented  in  Table  4. 


Table  4. — Amount  of  Water,  Freezing  at  -5°C.  in  Cabbage  Leaves  Hardened 
in  Coldframe  for  Different  Lengths  of  Time. 


Treatment 

Percent 

dry 

matter 

Percent 

water 

Percent 

total 

water 

frozen 

In  100  gran 
grams  water 
frozen 

as  of  tissue 
grams  water 
unfrozen 

Not  hardened  

9.91 

90.09 

82.1 

73.96 

16.13 

Hardened  2 days  .... 

13.20 

86.80 

75.3 

65.32 

21.48 

Hardened  4 days  .... 

13.90 

86.10 

62.8 

54.47 

31.63 

Hardened  9 days  .... 

14.00 

86.00 

58.7 

50.48 

35.62 

Hardened  14  days  . . . 

14.79 

85.21 

54.6 

46.52 

38.69 

Hardened  16  days  ... 

18.7 

81.3 

51.0 

41.46 

39.84 

Hardened  20  days  . . . 

19.35 

80.65 

47.9 

38.63 

42.02 

It  appears  that  the  percentage  of  the  total  water  frozen  at  -5°C. 
decreases  as  the  plant  tissue  increases  in  hardiness.  At  the  same 
time  the  amount  of  total  moisture  in  the  plants  decreases,  accom- 
panied by  an  increasing  percentage  of  dry  matter.  The  relation  of 
degree  of  hardening  to  the  percentage  of  freezable  water  and  to  dry 
matter  content  is  shown  graphically  in  Figure  4.  The  dates  of  de- 
crease in  percentage  of  water  frozen  and  of  increase  in  percentage 
of  dry  matter  proceed  quite  rapidly  the  first  four  or  five  days  the 
plants  are  exposed  to  hardening  in  tli e coldframe.  After  this,  these 
changes  proceed  slowly  for  some  days  longer.  On  the  whole,  it  seems 
that  there  is  a close  correlation  between  the  degree  of  hardiness  and 
the  percentage  of  total  water  retained  in  the  unfrozen  condition. 
The  actual  amount  of  ice  per  gram  of  fresh  leaf  tissue  also  decreases 


46 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


with  the  degree  of  hardening,  while  the  actual  amount  of  water  re- 
maining unfrozen  increases  as  shown  in  Figure  5. 


The  Hardening  Process  in  Vegetable  Plants. 


47 


Influence  of  time  of  day  on  percentage  of  water  frozen. — McCool 
and  Millar80  found  that  the  time  of  day  influenced  both  the  depres- 
sion of  the  freezing  point  and  the  amount  of  water  frozen  at  a given 
temperature.  Their  experiments  with  various  cereal  plants  showed 
that  the  freezing  point  depression  of  the  leaves  increases  during  the 
forenoon,  declines  slightly  in  the  afternoon  and  almost  reaches  tho 
early  morning  value  by  midnight.  Over  the  same  period  the  per- 
centage of  total  moisture  varied  inversely  with  the  depression  of  the 
freezing  point,  but  to  a much  less  degree.  Shaded  oat  plants  de- 
creased steadily  in  sap  density  during  the  day,  while  exposed  plants 
showed  the  usual  increase  at  mid-day.  The  slight  difference  in 
water  content  of  plants  is  held  by  these  writers  to  be  insufficient  to 
explain  fully  the  increased  sap  concentration  at  mid-day,  hence 
it  seems  that  the  products  of  photosynthesis  must  play  a part.  Bar- 
ley plants  kept  under  bell  jars  in  a saturated  atmosphere,  under  con- 
ditions retarding  transpiration  but  permitting  photosynthesis,  had  55> 
percent  of  the  water  in  the  tops  and  62  percent  of  that  in  the  roots 
frozen  at  -3°C.  to  -4°C.  in  the  morning.  At  noon  43  percent  of  the 
water  was  frozen  in  the  tops  and  59  percent  in  the  roots,  at  the  same 
temperature. 

It  has  been  shown  by  Dixon133  that  illumination  increased  the 
osmotic  concentration  in  leaves  and  this  concentration  gradually  fell 
when  light  was  cut  off.  Chandler20  also  found  that  plants  shaded  24 
hours  had  decreased  concentration.  According  to  Drabble  and 
Drabble,134  a greater  concentration  of  cell  sap  occurs  in  plants  sub- 
jected to  factors  favoring  rapid  loss  of  water  by  transpiration.  Under 
these  conditions  the  increased  concentration  of  cell  sap  is  probably 
very  largely  the  result  of,  as  well  as  the  means  of  protection  against, 
rapid  loss  of  water  from  the  leaves. 

In  the  course  of  the  experiments  on  the  amount  of  water  freezing 
in  cabbage  leaves  of  different  degrees  of  hardiness,  some  data  were 
obtained  relative  to  the  effect  of  the  time  of  day  on  the  amount  of 
water  freezing  in  leaves  of  the  same  hardiness.  No  attempt  was 
made  to  provide  specially  controlled  conditions ; they  were  the  same 
as  those  previously  described  for  the  various  hardening  treatments, 
and  were  identical  with  those  referred  to  in  Table  2. 


48 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Table  5. — Effect  of  Time  of  Day  on  Amount  of  Water  Frozen  in  Cabbage 

Leaves  at  -5°C. 


Material 

Time 

Percent  moisture 
in  plants 

Percent  water  frozen 
at  — 5'  C. 

Wet-grown  greenhouse 

9 A.  M. 

90.43 

82.4 

plants  (tender) 

2 P.  M. 

90.22 

78.2 

6 P.  M. 

85.9 

Dry-grown  greenhouse 

9 A.  M. 

86.60 

55.8 

plants  (hardy) 

2 P.  M. 

86.39 

47.1 

Coldframe 

9 A.  M. 

87.87 

61.9 

hardened  plants 

2 P.  M. 

84.12 

55.5 

It  is  seen  that  the  amount  of  freezable  water  is  somewhat  greater 
in  the  morning  than  in  early  afternoon,  but  the  differences  are  not 
as  great  as  those  found  by  McCool  and  Millar.  The  moisture  con- 
tent is  also  somewhat  less  in  the  afternoon  indicating  the  possibility 
of  a greater  power  of  imbibition  at  that  time.  Probably  the  larger 
factor  in  causing  the  slight  difference  in  amount  of  frozen  water  is 
the  increased  concentration  of  sugars  formed  by  the  photosynthetic 
activities  of  the  leaf.  Both  the  moisture  content  and  the  concentra- 
tion of  cell  sap  evidently  have  some  influence  on  the  amount  of 
freezable  water. 

Effect  of  watering  plants  with  salt  solutions  on  amount  of 
easily  frozen  water  in  the  leaves. — A method  used  to  harden  vege- 
table plants  was  watering  with  salt  solutions.  Only  one  of  these  ex- 
periments will  be  discussed  here.  On  February  15,  seedling  cabbage 
plants  were  potted  in  3-inch  clay  pots,  which  were  plunged  in  soil 
on  a raised  bench  in  the  greenhouse.  One,  series  was  potted  in  river 
sand,  one  in  greenhouse  compost  soil  and  a third  in  compost  plus 
rotten  stable  manure.  Each  series  was  divided  into  four  plots  and 
after  the  plants  were  well  established  one  of  each  series  was  watered 
with:  (1)  tap  water,  (2)  M/10  NaN03,  (3)  M/10  KC1,  (4)  M/10 
NaCl.  These  applications  were  repeated  every  few  days,  when  water 
appeared  to  be  required.  After  the  second  application,  the  rate  of 
growth  in  the  different  plots  was  evidently  being  affected.  All  of 
the  salt  solutions  depressed  growth  but  particularly  in  the  series 
grown  in  compost  and  in  the  compost  and  manure  mixture.  Plants 
growing  in  the  sand  showed  some  of  this  stunting  effect,  but  much 
later  than  in  compost  soils.  A test  made  March  30  showed  that  the 
plants  grown  in  the  compost  soils  and  stunted  by  the  salt  applica- 
tions were  much  hardier  than  those  receiving  tap  water  and  making 
normal  growth.  Little  effect  of  the  salts  upon  either  the  size  or 


The  Hardening  Process  in  Vegetable  Plants. 


49 


hardiness  of  the  plants  grown  in  sand  could  be  observed.  Plants 
grown  in  the  compost  soils  and  watered  with  NaCl  were  exposed  to 
-6°C.  for  45  minutes  without  injury.  Plants  in  compost  soils  watered 
with  KC1  and  NaN03  were  injured  somewhat  under  the  same  con- 
ditions, and  those  receiving  tap  water  were  killed.  Plants  from  all 
of  the  lots  grown  in  sand  were  killed  at  -6°C.,  but  when  exposed 
to  -3°C.  to  -4°C.  for  one  hour,  only  those  receiving  water  were  much 
injured.  The  day  following  the  freezing  tests,  dilatometer  determina- 
tions were  made  on  leaves  of  plants  from  some  of  the  lots  given 
different  treatments,  the  results  being  shown  in  Table  6.  Most  of 
these  figures  represent  only  one  determination.  The  samples  were 
gathered  about  1 :30  P.  M.  on  a bright  sunny  day,  which  may  explain 
why  the  percentage  of  water  frozen  in  some  of  these  plants  is  a little 
less  than  that  shown  in  Table  3 for  tissues  of  approximately  the 
same  degree  of  hardiness. 


Table  6. — Amount  of  Water  Freezing  at  -5°C.  in  Cabbage  Leaves  From 
Plants  Watered  with  Various  Salt  Solutions. 


iPercent  dry 
matter 

Treatment  of  plants 

Percent 

moisture 

Percent  total 
water  frozen 
at  -5°C. 

Grams 
water 
frozen  in 
100  grams 
of  leaves 

In  compost  soil  watered  with 
tap  water  (medium  tender)  . . 

.10.86 

89.14 

61.2 

54.55 

In  compost  soil  watered  with 
*1/10  NaN03  (hardy)  

. .11.83 

88.13 

37.3 

32.87 

In  compost  soil  watered  with 
M/10  NaCl  (hardy)  

.12.02 

87.98 

39.5 

34.76 

In  compost  and  manure-watered 
with  M/10  NaCl  (very  hardy) 

14.03 

85.97 

27.2 

23.29 

In  sand  watered  with 
tap  water  (very  tender)  

. 8.24 

91.76 

79.8 

73.22 

In  sand  watered  with 
M/10  NaCl  (medium  tender) 

11.01 

88.99 

59.4 

52.86 

The  percentage  of  water  frozen  is  much  less  in  the  stunted 
plants — those  found  most  hardy  to  cold.  The  amount  of  water  frozen 
is  correlated  with  the  observed  degree  of  cold-resistance  and  the 
extent  to  which  growth  was  checked.  Here  again  the  percentage  of 
water  frozen  varies  inversely  with  the  percentage  of  dry  matter. 
Unfortunately  the  freezing  point  depressions  of  the  plants  used  in 
this  experiment  were  not  taken.  However,  we  know  from  Chandler’s 
work  that  the  sap  of  plants  watered  with  salt  solutions  has  an  in- 
creased osmotic  concentration.  Bartetzko4  found  that  Aspergillus , 
Penicillium  and  other  fungi  grown  in  nutrient  media  of  varying 


50 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


concentrations  increased  their  resistance  to  freezing  in  proportion  to 
the  increase  in  the  osmotic  strength  of  the  medium. 

The  question  arises  as  to  how  the  application  of  salt  solutions 
to  soils  in  which  plants  are  growing  checks  growth,  increases  cold 
resistance  and  reduces  the  amount  of  freezable  water.  The  stunt- 
ing might  be  due  to:  (a)  The  toxicity  of  the  salt  solution  to  the 
roots  of  the  plants  at  the  concentration  used.  However,  since  the 
salt  solutions  were  not  appreciably  toxic  to  the  plants  grown  in  sand, 
it  seems  doubtful  if  the  stunting  and  hardening  of  the  cabbage  plants 
in  the  compost  soils  can  be  attributed  to  this  factor,  (b)  Absorption 
of  the  salts  by  the  plants,  causing  a greater  concentration  of  the  sap, 
yet  why  should  nutrient  salts  such  as  NaN03  cause  a stunting  of 
healthy  plants?  (c)  Condition  of  physiological  drought  within  the 
plant,  at  least  at  such  times  as  the  moisture  content  of  the  soil  was 
low  or  the  rate -of  transpiration  very  rapid.  Such  a condition  might 
easily  arise,  in  treating  a succulent  plant  such  as  cabbage,  with  rather 
strong  salt  solutions.  If  the  tops  of  the  plants  suffered  from  phy- 
siological drought  a considerable  part  of  the  time  because  the  roots 
were  unable  to  absorb  water  rapidly  from  the  more  concentrated  soil 
solution,  then  a condition  would  exist  more  or  less  similar  to  that 
in  ordinary  soils  wherein  plants  have  been  hardened  by  partially 
withholding  moisture.  The  fact  that  the  lots  grown  in  sand  did  not 
show  nearly  so  much  of  the  checking  or  stunting  effect  as  those 
grown  in  the  finer  soils  containing  more  organic  matter  lends  strength 
to  this  idea.  To  see  whether  or  not  the  observed  results  might  be 
due  to  variations  in  the  concentration  of  the  soil  solution  this  was 
determined  in  each  lot  at  the  end  of  the  experiment.  The  method  of 
Bouyoucos13  wras  employed,  taking  15  grams  of  air-dry  soil  and  10 
cc.  of  distilled  water,  determining  the  freezing  point  depression  with 
the  Beckman  thermometer,  and  calculating  that  the  soil  solution 
contained  100  parts  of  solute  per  million  for  each  0.004° C.  of  freezing 
point  lowering.  The  results  are  presented  in  Table  7. 


The  Hardening  Process  in  Vegetable  Plants. 


51 


Table  7. — Concentration  of  Soil  Solutions  After  M/10  Salt  Solutions 
Were  Applied  Five  Times. 


Treatment 

Sand 

Compost 

Compost  and  manure 

Freezing  1 
point 

depression 

p.p.m.  in 
soil 

solution 

Freezing 

point 

depression 

p.p.m.  in 
soil 

solution 

Freezing 

point 

depression 

p.p.m.  in 
soil 

solution 

Tap  water  . . 

.005°C. 

125 

.045°C. 

1125 

.159°C. 

3975 

M/10  NaNOs  . . 

.020°C. 

400 

.277 

6925 

M/10  KC1  .. 

.020°C. 

400 

.210 

5200 

M/10  NaCl  . . 

1 .020°C. 

400 

.263 

6576 

: .367 

9175 

The  results  set  forth  in  Table  7 show  that  at  the  conclusion  of 
the  experiment,  just  after  samples  for  the  dilatometer  determinations 
had  been  taken,  the  concentration  of  the  soil  solutions  had  increased 
very  markedly  in  the  compost  soils  to  which  the  salts  were  applied, 
as  compared  to  the  check  of  the  same  sort  of  soil,  but  receiving  tap 
water.  However,  in  sand  the  soil  solution  was  much  less  concentrated 
and  there  was  no  great  increase  in  the  concentration  of  the  soil 
solutions  where  the  salts  were  applied.  Bouyoucos13  has  shown  that 


52 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


soils  containing  much  organic  matter  cause  a large  amount  of  prac- 
tically pure  water  to  become  “unfree”  by  means  of  capillary  ad- 
sorption. At  a given  moisture  content  therefore,  the  free  solution 
in  such  soils  would  be  more  concentrated  than  in  a sandy  soil  having 
little  organic  matter  and  large  soil  particles.  Comparing  the  data  in 
Tables  6 and  7 it  is  observed  that  the  amount  of  water  frozen  in 
plants  at  -5°C.  varies  inversely  to  the  concentration  of  the  soil  solu- 
tion, but  probably  not  in  proportion  to  it.  This  point  is  illustrated 
by  Figure  6. 

To  show  to  what  extent  the  growth  of  plants  in  this  experiment 
was  affected  by  the  salt  applications  in  the  three  soil  media  used,  the 
average  green  weights,  average  dry  weights,  and  percentages  of  dry 
matter  are  given  in  Table  8,  for  the  plants  in  each  lot  at  the  end 
of  the  experiment  (one  day  after  the  freezing  determinations  were 
made). 


Table  8. — Average  Growth  Made  by  Cabbage  Plants  in  Soils  Receiving 
Five  Applications  of  M/10  Salt  Solutions. 


Treatment 

Sand 

Sandy  compost  J 

Sandy  compost  and 
horse  manure 

Green 

Dry 

% dry 

Green 

Dry 

% dry 

Green 

dry  1 

% dry 

wt. 

wt. 

matter 

wt. 

wt. 

matter 

wt. 

wt. ! 

matter 

Tap  water  . . . 

13.55 

.967 

6.78 

8.12 

.581 

7.16 

M/10  NaN03  . 

8.58 

.560 

6.52 

4.63 

.431 

9.31 

M/10  KC1  . . 

8.32 

.523 

6.29 

5.38 

.505 

9.40 

6.18 

.631 

; 10.22 

M/10  NaCl  . . 

8.56 

.546 

| 6.37 

4.76 

.455 

9.57 

5.53 

1 .624 

j 9.55 

It  is  seen  from  Table  8 that  the  plants  in  the  sand  made  nearly 
twice  as  much  growth  as  plants  receiving  corresponding  treatment 
in  the  compost  soils,  using  the  average  green  weights  as  the  indicator. 
It  should  be  remembered  that  in  this  experiment  a rather  high  soil 
moisture  was  maintained  in  order  to  prevent  the  lack  of  water  from 
influencing  the  results ; furthermore  this  experiment  was  ended 
before  the  lack  of  nutrient  material  in  the  sandy  soils  could  become 
the  main  factor  limiting  their  growth. 

The  indications  point  to  the  conclusion  that  applications  of 
rather  strong  salt  solutions  raised  the  concentration  of  the  soil  solu- 
tion to  a point  at  which  roots  could  take  up  water  only  slowly,  and 
probably  not  at  all  when  the  total  soil  moisture  content  fell  below  a 
certain  point.  This  developed  a state  of  physiological  drought  in  the 
tops  due  to  the  restricted  water  intake.  Under  these  conditions,  the 
leaves  developed  xerophytic  characteristics  to  some  extent,  as  indi- 


The  Hardening  Process  in  Vegetable  Plants. 


53 


cated  by  the  greatly  increased  water-retaining  power  on  the  part  of 
the  cells.  This  is  shown  by  the  smaller  amounts  of  water  frozen  in 
the  leaves  of  such  plants,  and  as  is  shown  later,  by  the  lower  trans- 
piration rate,  and  slower  rate  of  drying  in  an  oven.  Another  ex- 
ample of  increased  cold  resistance  apparently  resulting  from  phy- 
siological drought  was  observed  in  the  field  in  the  spring  of  1921. 
On  March  31,  the  temperature  fell  to  -8°C.,  and  the  following  night 
to  -6°C.  Hardened  cabbage  plants  set  in  the  field  10  days  previous, 
were  very  severely  injured,  but  here  and  there  thru  the  field  small 
plants  were  observed  after  the  freeze,  the  leaves  of  which  were  ap- 
parently uninjured.  On  examination,  the  stems  of  all  such  plants 
were  found  to  be  nearly  severed  by  a “ damping  off”  fungus.  Evi- 
dently the  stem  injury  by  the  fungus  had  caused  physiological  drought 
in  the  top  of  the  plant,  resulting  in  considerable  increase  in  hardiness. 

Relation  of  amount  of  freezable  water  to  percentage  of  dry 
matter  and  freezing  point  depression  in  garden  plants. — Three  spec- 
ies of  plants  were  used  in  these  experiments,  cauliflower  representing 
a group-  possessing  potential  hardiness,  and  tomatoes  and  sweet  po- 
tatoes representing  plants  lacking  potential  hardiness.  Leaves  were 
gathered  during  June  from  plants  growing  under  ordinary  condi- 
tions in  the  garden.  The  soil  was  fairly  moist  at  this  time  and  the 
plants  were  making  good  growth.  A portion  of  each  lot  of  leaves 
was  used  for  the  dilatometer  determination,  and  another  portion  for 
determination  of  the  freezing  point  depression.  This  latter  was  made, 
not  on  the  expressed  sap,  as  were  the  previous  determinations  herein 
reported,  but  directly  on  the  triturated  leaf  tissue,  according  to  the 
method  of  Bouyoucos  and  McCool.14  The  results  are  given  for  two 
sets  of  determinations  in  Table  9.  In  the  last  two  columns  of  this 
table  are  given  the  relative  amounts  of  frozen  and  unfrozen  water, 
calculated  on  the  basis  of  100  grams  of  fresh  leaf  tissue. 

The  plants  used  in  these  experiments  would  probably 
have  been  killed  by  a brief  exposure  to  -3°C.,  except  the  cauli- 
flower, which  might  have  withstood  a somewhat  lower  temperature. 
It  may  be  seen  from  Table  9 that  the  percentage  of  total  water 
freezing  in  cauliflower  at  -5°C.  is  somewhat  less  than  in  tomato  and 
sweet  potato,  while  at  -3°C.  this  difference  is  much  greater  in  favor 
of  the  hardier  cauliflower.  The  amount  of  water  remaining  unfrozen 
is  correspondingly  greater  in  cauliflower.  It  appears  that  allowing 
cauliflower  leaves  to  stand  in  8 percent  sucrose  over  night  has  in- 
creased the  percentage  of  dry  matter  and  the  freezing  point  depres- 
sion and  has  decreased  the  amount  of  water  freezing  at  -5°C.  The 


54 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Table  9. — Amount  of  Water  Frozen  in  Leaves  of  Garden  Plants. 


In  100  grams  leaf  tissue 

frozen  at  - 

-5°C. 

% 

Freezing 

grams 

grams 

dry 

point 

% water 

water 

water 

Date 

Plant 

matter 

depression 

frozen 

frozen 

unfrozen 

June  8 

Cauliflower 
(in  8%  sucrose 

15.86 

.780°C. 

57.0 

48.0 

36.14 

over  night) 
Cauliflower 

13.26 

.413 

77.4 

67.1 

19.64 

(in  water  over 
night) 

Tomato 

11.73 

.650 

79.3 

70.0 

18.27 

99  99 

Sweet  Potato 

17.5 

83.0 

69.4 

13.1 

Frozen  at  - 

-3°C. 

June  21 

Cauliflower 

17.7 

1.300 

28.3 

23.3 

59.0 

" " 

Tomato 

13.5 

.915 

43.7  | 

37.9 

48.6 

" " 

Sweet  Potato 

14.84 

.750 

48.1 

41.0 

44.16 

amount  of  water  remaining  unfrozen  in  the  cauliflower  leaves  which 
had  been  in  the  sugar  solution  is  nearly  twice  as  much  as  in  the 
check  leaves  kept  in  water.  Since  sucrose  penetrates  plant  tissue 
quite  slowly,  the  changes  noted  are  probably  not  due  to  increased 
sugar  content.  However,  the  dry  matter  is  2.60  percent  or  nearly 
J greater  in  the  leaves  placed  in  sugar  solution.  The  8 percent 
sucrose  solution  is  approximately  equivalent  to  0.25  molecular  con- 
centration. Under  these  conditions,  water  may  be  withdrawn  from 
the  leaf,  thereby  decreasing  the  moisture  content,  increasing  the  per- 
centage of  dry  matter  and  presumably  increasing  the  power  of  im- 
bibition with  which  the  remaining  moisture  is  held  by  the  leaf  cells. 
Rather  tender  cabbage  plants,  the  roots  of  which  were  placed  in  8 per- 
cent sucrose  solution,  wilted  quickly,  indicating  withdrawal  of  water 
from  the  upper  portion  of  the  plant,  or  at  least  stoppage  of  intake 
to  make  up  losses  by  transpiration. 

Resume.— It  does  not  necessarily  follow  from  the  water-loss 
theory  of  killing  by  cold  that  there  is  a definite  minimum  moisture 
content  below  which  the  protoplasm  of  all  plants  dies.  In  view  of 
experiments  such  as  those  of  Adams3  and  of  Kiesselbach  and  Rat- 
cliff52 it  seems  quite  likely  that  the  minimum  amount  of  water  re- 
quired by  plant  cells  to  retain  life  varies  with  the  state  of  physiologi- 
cal activity,  the  stage  of  development,  perhaps  with  changes  in  either 
internal  or  external  conditions,  and  probably  differs  in  various  spe- 
cies at  the  same  stage  of  development  and  under  the  same  conditions. 
Ewart135  has  shown  that  some  seeds  can  he  dried  to  a moisture  con- 


The  Hardening  Process  in  Vegetable  Plants. 


55 


tent  of  1 or  2 percent  without  killing  and  there  is  reason  to  believe 
that  if  a tender  cabbage  leaf  is  killed  by  the  loss  of  50  percent  of  its 
water,  a hardened  leaf  may  be  able  to  survive  the  loss  of  even  a 
larger  fraction  at  still  lower  temperatures.  May  not  the  hardening 
process  in  vegetable  plants,  the  maturing  process  in  woody  stems 
and  the  ripening  process  in  seeds  involve  changes  which  increase  the 
stability  of  the  protoplasmic  structure  as  well  as  changes  which  make 
for  increased  water-retaining  power? 

RATE  OF  WATER-LOSS  BY  TRANSPIRATION  IN  HARDENED 
AND  TENDER  CABBAGE. 

It  is  a commonly  observed  fact  that  non-hardened  vegetable 
plants  wiiLseverely.  upon  transplanting  to  the  field  and  if  conditions 
favor  rapid  transpiration  or  if  the  soil  is  dry  they  may  die,  due  to 
excessive  water  loss.  On  the  other  hand,  plants  properly  hardened 
by  any  of  the  methods  mentioned  in  this  paper  withstand  transplant- 
ing without  serious  wilting.  To  the  practical  grower  the  ability  of 
hardened  plants  to  survive  transplanting  without  dangerous  wilting 
is  probably  of  greater  importance  than  the  increased  cold-resistance 
developed  by  the  hardening  process.  Plate  6,  B and  C,  illustrates  the 
marked  difference  in  turgor  of  hardened  and  not  hardened  cabbage 
plants  one  day  after  transplanting  to  the  field.  These  were  potted 
plants,  so  the  root  systems  were  not  disturbed  much  by  transplanting. 

Of  interest  in  this  connection  are  the  observations  of  Bergen7 
on  the  rate  of  transpiration  of  a number  of  evergreens,  as  Olea , Qiter- 
cus  and  Pistacia,  compared  to  that  of  TJlmus  and  Pisum  sativum.  He 
found  that  the  water  loss  in  the  former  group  was  25  percent  less 
than  in  the  latter.  He  concluded,  however,  that  xerophytic  leaf  struc- 
ture (of  the  hardy  evergreens)  is  not  always  incompatible  with 
abundant  transpiration,  but  sometimes  exists  only  for  use  in  emer- 
gencies, to  protect  the  plant  from  injurious  loss  of  water. 

Salmon108  draws  attention  to  the  xerophytic  structure  of  the 
hardiest  types  of  winter  cereals;  winter  rye,  Turkey  and  Kharkoff 
wheats  are  characterized,  for  example,  by  a narrow  leaf  and  pros- 
trate habit  of  growth.  The  same  is  true  of  Winter  Turf,  the  hardiest 
variety  of  winter  oats.  Salmon  found  no  differences  in  cell  structure, 
epidermal  covering,  or  mechanical  ability  to  control  transpiration, 
that  could  be  correlated  with  the  great  difference  in  hardiness  known 
to  exist  in  cereals,  except  that  Turkey  wheat  (hardy)  had  25  percent 
greater  root  length  than  Fultz  (less  hardy)  and  40  percent  greater 
than  oats  and  barley  (least  hardy).  This  character  might  enable  the 


56 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


plant  to  escape  dangerous  drying  out  when  the  ground  is  frozen  to 
a certain  depth. 

The  relation  existing  between  water-retaining  power  and  re- 
sistance to  cold  is  demonstrated  by  observations  of  workers53  in  the 
United  States  Forest  Service,  in  a recent  study  of  a chlorosis  of  coni- 
fer seedings.  The  chlorotic  leaves  were  less  turgid  than  normal 
leaves  and  wilted  very  quickly  when  the  water  supply  was  cut  off; 
in  fact,  chlorotic  leaves  of  the  Douglass  fir  wilted  so  quickly  that 
accurate  leaf  measurements  could  not  be  made.  Plants  having  chlo- 
rotic leaves  failed  to  harden  properly  in  the  fall,  so  that  many  were 
injured  by  early  fall  frosts  and  many  more  by  winter  cold.  How- 
ever, in  plots  where  the  chlorosis  was  corrected  in  summer  by  spray- 
ing with  ferrous  sulfate,  the  plants  became  perfectly  winter-hardy. 
Evidently  chlorotic  leaves  are  unable,  because  of  absence  of  chlo- 
rophyl,  to  develop  the  usual  water-retaining  power  and  cold  resistance 
of  the  species. 

It  was  considered  desirable  to  determine  the  difference  in  rate 
of  transpiration  of  non-hardened  plants  and  plants  hardened  in 
various  ways,  because  of  the  indications  which  might  be  obtained 
thereby  as  to  the  relative  water-retaining  power  of  plants  of  different 
degrees  of  hardiness.  Four  experiments  were  performed,  using  cab- 
bage plants  in  4-inch  clay  pots.  The  pots  were  coated  and  sealed 
with  a mixture  of  paraffin,  vaseline  and  beeswax.  Two  to  four  plants 
were  used  from  each  experimental  lot.  Before  sealing,  the  pots  were 
brought  to  uniform  moisture  content.  The  experiments  were  con- 
ducted under  different  conditions,  but  in  each  experiment  the  plants 
were  kept  uniform  with  reference  to  external  factors.  Plants  as 
nearly  the  same  size  as  possible  were  used,  but  the  hardened  plants 
were  usually  smaller  than  the  non-hardened.  At  the  conclusion  of 
each  experiment  the  plants  were  weighed  at  once  and  the  leaf  area 
of  each  plant  was  measured  with  a planimeter.  The  results  of  the 
four  experiments  are  presented  in  Table  10. 


The  Hardening  Process  in  Vegetable  Plants. 


57 


Table  10. — Transpiration  Experiments  With  Cabbage  Plants. 


Treatment  of  plants 

No. 

plants  j 
used  | 

Av.  leaf 
area  per 
plant 

Av.  Ajmit. 
transpired 
per  plant  , 

Transpiration  in  grains 
per  hour 

j per  sq.  M. 
per  plant  ! leaf  area 

Expt.  1 (outdoors)  3/15/21  partly  cloudy, 

cool,  moderate  wind, 

24  hours 

Dry-grown  gh.  plants 

4 

125.0  sq.  cm. 

13.15  g. 

0.547 

43.7 

Wet-grown  gh.  plants 

2 

354.0 

39.6 

1.649 

46.6 

Expt.  2 (In  cool  greenhouse)  3/15/21  temp. 

60-70  degrees  F.,  24 

hours 

Dry-grown  gh.  plants 

4 

167.0 

12.95 

I 

0.539  i 

32.3 

Wet-grown  gh.  plants 

2 

285.0 

27.9 

1.162 

40.8 

Expt.  3 (In  warm  greenhouse)  3/19/21  temp.  65-80 

degrees  F.,  24  hours 

Coldframe  hardened 

1 

for  5 days  

2 

347.0 

34.60 

1.441 

41.5 

Dry-grown  gh.  plants 

4 ; 

165.0 

19.22 

0.800 

48.5 

Wet-grown  gh.  plants 

2 ! 

1 

315.0 

I ! 

41.15 

1.714 

54.4 

Expt.  4 (Outdoors)  4/2/21,  clear,  warm,  little  wind, 

, 5 hours,  11:30  A.  M. 

to  4:30  P.  M. 

Coldframe  hardened 

for  1 week  

2 

278.3 

i 18.90 

3.778 

135.6 

Med.  dry-grown  gh. 

plants  

3 

202.0 

12.97 

2.590 

128.3 

Med.  wet-grown  gh. 

plants  

2 

395.5 

1 29.6 

5.920 

150.0 

Greenhouse  plants 

grown  in  compost  soil 

and  watered  with  M/10 

NaCl  (hardy) 

2 

153.6 

j 10.5 

2.100 

136.6 

Same,  watered  with 

tap  water  (tender)  ...2 

164.0 

15.9 

3.180 

193.9 

Greenhouse  plants 

grown  in  sand,  and 

watered  with  tap 

water  (tender)  . 

2 

| 252.8 

1 23.9 

4.780 

189.0 

The  water  loss  per  square  centimeter  of  leaf  area  per  hour  is 
somewhat  greater  in  tender  plants  than  in  those  hardened  by  drying, 
by  coldframe  exposure,  or  by  watering  with  salt  solutions.  The 
much  greater  total  water  loss  of  the  non-hardened  plants  was  due 
to  a large  extent  to  the  fact  that  they  were  larger  than  the  hardened 
plants,  though  of  the  same  age. 


58 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


The  fact  that  the  rate  of  transpiration  per  unit  of  leaf  area  was 
less  in  hardened  plants  is  significant.  If  the  rate  of  diffusion  of 
water  from  the  cells  into  the  intercellular  spaces  determines  the  rate 
of  transpiration,  then  a lower  rate  of  transpiration  would  be  as- 
sociated with  a greater  water-retaining  power  on  the  part  of  the 
plant  cells.  This  water-retaining  power  would  be  exerted  when  the 
plant’s  cells  are  exposed  to  water  loss  by  freezing  in  the  same  way 
as  when  exposed  to  loss  by  transpiration  or  by  drying. 

RATE  OF  DEHYDRATION  IN  HARDENED  AND  TENDER 

PLANTS. 

Since  it  was  found  that  hardened  plants  exhibited  a greater 
water-retaining  power  than  non-hardened  plants  upon  freezing,  it 
was  thought  that  the  difference  might  be  measured  by  the  rate  of 
water  loss  in  similar  tissues  exposed  to  drying. 

Mr.  V.  R.  Boswell8  undertook  a special  investigation  of  the  rate 
of  dehydration  of  leaves  from  hardened  and  non-hardened  plants 
during  the  winter  and  spring  of  1921.  The  material  used  in  his 
experiments  was  from  the  same  lots  upon  which  other  results  are 
reported  in  this  paper. 

Leaves  of  uniform  condition  and  from  corresponding  parts  of 
plants  were  gathered  from  cabbage  and  tomato  plants  subjected  to 
various  hardening  treatments.  Lots  directly  comparable  were  gath- 
ered and  dried  at  the  same  time.  The  samples  were  placed  in  stop- 
pered bottles,  taken  at  once  to  the  laboratory,  weighed,  and  immediate- 


Table  11. — Rate  of  Water  Loss  by  Drying  at  60°C.  in  Hardened  and  Ten- 
der Leaveis. 

(In  per  cent  of  total  moisture  content) 


Tomato 

leaves 

Cabbage  leaves 

Time 

Greenhouse 

Hardened 

Greenhouse 

Hardened 

Greenhouse  plants 

in 

wet-grown 

in  cold- 

wet-grown 

in  cold- 

water 

NaNO3  | NaCl 

minutes 

(tender) 

frame 

(tender) 

frame 

(ten- 

(medi- (hardy) 

der) 

um  har-  , 

dy)  | 

15 

34.77 

26.46 

21.53 

8.92 

23.82 

19.70  ’ 12.13 

30 

68.11 

57.91 

42.71 

17.43 

46.71 

38.68  24.29 

45 

83.99 

75.34 

54.20 

36.83 

62.74 

54.80  34.68 

60 

94.27 

87.85 

75.32 

47.56 

74.81 

j 66.27  43.13 

75 

97.94 

95.57 

79.08 

55.23 

84.67 

| 76.42  51.99 

90 

99.34 

99.02 

93.32 

68.23 

91.59 

| 83.20  59.36 

105 

99.59 

99.52 

93.12 

74.58 

96.32 

! 89.27  66.79 

120 

99.62 

99.61 

99.66 

86.01 

98.73 

! 93.94  73.34 

135 

99.64 

99.63  | 

98.11 

94.88 

99.68 

j 97.57  i 78.87 

150 

99.80 

94.73 

99.93 

! 99.37  84.48 

165 



1 

98.88 

99.95 

1 99.87  88.91 

The  Hardening  Process  in  Vegetable  Plants. 


59 


ly  placed  in  an  electric  oven  at  a constant  temperature  of  60°C.  The 
leaves  were  spread  out  on  wire  gauze  placed  on  the  shelves  of  the 
oven  until  they  were  beginning  to  become  brittle,  after  which  they 
were  transferred  to  the  weighing  bottles  in  which  the  dehydration 
was  completed.  Each  lot  of  leaves  was  removed  from  the  oven  at 
intervals  of  15  minutes,  cooled  and  weighed.  From  the  loss  in  weight 
for  each  period  of  drying,  was  calculated  the  percent  of  total  mois- 
ture removed  per  period.  Table  11  compiled  from  some  of  Boswell’s 
data,  gives  the  percent  of  total  moisture  lost  at  the  end  of  each  15- 
minutes  interval  in  samples  of  hardened  and  tender  plants. 

The  data  presented  in  Table  11  bring  out  a striking  difference 
in  the  rate  of  water-loss  by  drying  in  hardened  and  non-hardened 
leaves  of  cabbage,  especially  at  the  beginning  of  the  period  of  drying. 
This  difference  as  not  very  great  in  the  two  lots  of  leaves  of  tomato. 
In  cabbage,  leaves  from  plants  hardened  by  exposure  in  the  cold- 
frame,  by  watering  with  salt  solutions  and  by  partially  withholding 
water,  show  a much  smaller  loss  of  water  for  each  period  than  do 
leaves  of  tender,  well-watered  plants  grown  in  the  greenhouse.  This 
difference  in  rate  of  drying  indicates  a relatively  much  greater 
water-retaining  power  in  hardened  plants.  Whatever  the  differences 
in  the  two  types  of  plant  tissue  are,  the  greater  water-retaining  power 
of  the  hardy  tissue  evidently  does  not  depend  entirely  on  the  or- 
ganization of  living  matter,  but  on  the  chemical  and  physical  prop- 
erties of  the  substances  of  which  the  tissues  are  composed. 

Another  point  which  may  be  seen  from  Boswell’s  dehydration 
experiments  is  that  tomato  leaves  dry  out  much  more  rapidly  than 
cabbage  leaves.  Even  the  hardened  tomato  leaves  give  up  water 
faster  than  the  leaves  of  non-hardened  cabbage.  In  view  of  the  fact 
that  the  tomato  is  not  susceptible  of  hardening  to  the  extent  of  sur- 
viving ice  formation,  it  seems  that  we  have  here  an  indication  of 
the  fundamental  difference  between  the  two  types  of  plants.  The 
tomato  lacks  the  potential  ability  to  acquire  or  develop  increased 
water-retaining  power  to  any  great  degree,  while  the  cabbage  and 
similar  plants  have  this  potentiality  to  a considerable  degree. 


60 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Figure  7 shows  graphically  the  relative  rate  of  water-loss  by 
diying  in  the  different  types  of  tissue. 


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Fig.  7. — Rate  of  water  loss  from  leaves  of  varying  degrees  of  hardiness. 

CHANGES  IN  CARBOHYDRATES  ON  HARDENING  OF 

PLANTS. 

Formation  of  sugar  by  low  temperature.— Numerous  investi- 
gators have  noted  increased  amounts  of  sugars  in  plants  exposed  to 
low  temperature.  Mer67  was  probably  the  first  to  note  the  disap- 
pearance of  starch  and  the  accumulation  of  sugar  in  evergreen  leaves 
in  winter. 

Lidforss60  noted  in  a number  of  evergreen  plants  in  Sweden  that 
starch  was  converted  to  sugar  in  the  fall  and  reconverted  into  starch 
in  spring.  He  found  that  tender  seedlings  placed  in  a sugar  solu- 
tion for  a short  time  w^ere  able  to  withstand  several  degrees  of  lower 
temperature  without  injury.  Lidforss  thought  the  hardiness  of  the 
evergreen  leaves  and  the  sugar-treated  seedlings  wTas  due  to  increased 
concentration  of  the  cell  sap  resulting  from  the  accumulation  of 
sugar,  to  which  he  attributed  reduced  transpiration  and  lowrer  freez- 
ing point  depression,  as  well  as  a protective  effect  of  sugar  on  the 
precipitation  of  proteins  of  the  cell.  Gorke35  found  that  he  could 
prevent  the  precipitation  of  protein  from  expressed  plant  sap  by 
adding  sugar. 


The  Hardening  Process  in  Vegetable  Plants. 


61 


Miyake70  examined  the  leaves  of  evergreen  plants  in  various 
parts  of  Japan  in  winter  finding  those  of  many  plants  to  be  starch- 
free  during  the  coldest  part  of  winter.  Another  group  had  very 
little  starch  in  the  mesophyll  during  cold  weather  (when  the  mean 
temperature  was  near  or  below  the  freezing  point.)  Plants  in 
Northern  Japan  were  markedly  lower  in  starch  than  in  the  warmer 
sections.  Schulz111  examined  one  hundred  species  of  plants  in  Ger- 
many, finding  most  of  them  starch-free  in  winter,  while  a few  con- 
tained a little  starch  mostly  in  the  fibrovaseular  bundles  and  the 
surrounding  cells. 

Recently,  Swedish  investigators2  have  shown  that  the  hardier 
varieties  of  wheat  have  a larger  sugar  content  in  fall  and  winter. 
They  found  that  the  percentage  of  dry  matter  and  the  amount  of 
sugar  in  winter  wheat  varies  considerably  during  the  winter,  fluc- 
tuating with  the  temperature,  but  during  the  period  from  November 
12  to  February  15,  no  starch  could  be  found  in  the  leaves.  Gasner 
and  Grimme31  upon  analyzing  the  first  leaves  of  wheat,  found  that 
seedlings  germinated  at  5-6 °C.  had  a greater  sugar  content  than 
those  germinated  at  28 °C.  They  also  found  a higher  sugar  content 
in  leaves  of  hardy  winter  wheats  than  in  spring  wheats  germinated  at 
the  same  temperature. 

Micheal-Durand69  in  extensive  studies  on  the  changes  of  carbo- 
hydrates in  plants,  found  an  enormous  accumulation  of  sugars  in 
leaves  of  certain  evergreens  in  winter,  while  starch  completely  dis- 
appeared during  the  coldest  weather.  ITe  explains  this  condition  as 
follows : 

(1)  In  winter  assimilation  is  low,  but  respiration  is  depressed 
still  more  by  the  low  temperature. 

(2)  Conditions  in  winter  are  unfavorable  for  translocation. 

(3)  Low  temperature  prevents  the  condensation  of  the  simple 
sugars  into  higher  carbohydrates. 

(4)  Breaking  up  (splitting)  of  polysaccharides. 

Miiller-Thurgau75  and  others,  have  measured  the  accumulation 

of  simple  sugars  in  potatoes  at  the  expense  of  starch  upon  exposure 
to  low  temperature  and  the  reconversion  of  the  sugars  into  starch 
when  higher  temperatures  are  provided.  This  reversible  chemical 
change  seems  to  be  generally  associated  with  changing  temperatures 
near  the  freezing  point,  probably  due  to  shifting  of  chemical  equi- 
librium by  enzymatic  activity. 

Relation  of  sugar  content  to  cold  resistance.  — Since  the  ex- 
periments of  Lidforss00  and  Gorke35  the  extensive  formation  of 


62 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


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The  Hardening  Process  in  Vegetable  Plants. 


63 


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64 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


sugars  in  leaves  of  plants-  exposed  to  cold  has  generally  been  con- 
sidered to  be  related  to  their  cold-resistance.  However,  Harvey42 
concluded  that  carbohydrate  changes  were  not  important  in  the 
hardening  process  with  cabbage  plants,  since  he  found  that  cabbage 
plants  could  be  hardened  to  some  extent  at  least,  by  keeping  them 
several  days  in  the  dark  in  a low  temperature  chamber,  during  which 
time  there  was  little  change  in  the  carbohydrate  equilibrium.  How- 
ever, it  has  been  shown  by  several  investigators  and  notably  by  Lewis 
and  Tuttle59  that  simple  sugars  form  a large  part  of  the  osmotieally 
active  cell  contents. 

From  the  beginning  of  these  experiments,  samples  were  col- 
lected for  carbohydrate  analyses  from  some  of  the  series  of  plants 
in  each  of  the  hardening  treatments.  The  results  of  some  of  these 
determinations  are  given  in  Table  12. 

Methods  of  analysis. — The  sugar  analyses  were  made  according 
to  the  modified  Munsen  and  Walker  method,  as  described  by  Hook- 
er,46 the  results  being  expressed  as  dextrose. 


One  gram  of  the  air  dry,  ground  plant  material  was  weighed,  transferred 
to  filter  paper  and  washed  thoroughly  five  times  with  distilled  water.  The 
insoluble  residue  was  used  for  the  starch  determination.  The  filtrate, 
amounting  to  about  150  cc.,  was  taken  for  determination  of  soluble  sugars. 
After  clearing  with  basic  lead  acetate  the  extract  was  made  up  to  250  cc. 
and  filtered.  Two  hundred  cc.  of  the  filtrate  was  pipetted  into  a volumetric 
flask,  excess  lead  precipitated  with  solid  sodium  carbonate,  made  up  to 
250  cc.  and  filtered.  An  aliquot  of  the  filtrate  (Solution  A)  was  used  for 
the  determination  of  reducing  sugars,  while  another  portion  was  used  for 
determination  of  the  total  sugars. 

Five  cc.  of  concentrated  HC1  was  added  to  75  cc.  of  Solution 
A and  hydrolized  at  70°C.  for  exactly  ten  minutes  (Solution  B).  After 
cooling,  this  solution  was  neutralized  with  sodium  hydroxide  made  up  to 
100  cc.  and  used  for  the  determination  of  total  sugars  as  dextrose. 

The  sugar-free  residue  of  the  original  sample  was  used  for  the  starch 
determination.  It  was  washed  into  a beaker,  boiled  five  minutes  to  convert 
the  starch  into  a paste  and  after  cooling  3 cc.  of  Taka-diastase  solution  were 
added.  The  beaker  was  then  placed  in  the  oven  at  40°C.  for  24  hours, 
the  starch  being  broken  down  to  maltose  and  dextrin.  The  liquid  containing 
these  sugars  was  then  filtered  off,  adding  the  washings  to  the  filtrate,  which 
wTas  hydrolized  with  acid  for  2 y2  hours  under  a reflux  condenser  to  break 
down  further  the  products  of  digestion  to  dextrose.  After  cooling,  the  solu- 
tion was  neutralized  with  sodium  hydroxide,  cleared  and  prepared  for 
analysis  as  previously  described.  A blank  with  the  same  amount  of  Taka- 
diastase  solution  was  run  with  each  series  of  starch  determination. 

Total  polysaccharides  were  determined  on  a sample  of  the  dry  plant 
material  washed  free  of  soluble  sugars  with  cold  water.  The  filter  paper 
was  punctured  and  the  residue  washed  into  a 700  cc.  flask.  Eight  cc. 
concentrated  HC1.  and  enough  water  was  added  to  bring  the  total  volume 
to  150  cc.  After  boiling  two  and  one  half  hours  under  reflux  condenser, 
the  contents  of  flask  were  cooled,  transferred  to  a beaker  and  made  neutral 
to  litmus  with  sodium  hydroxide.  The  solution  was  then  prepared  for 
analysis  as  previously  described  for  Solution  A. 


The  Hardening  Process  in  Vegetable  Plants. 


65 


Discussion. — Table  12  presents  evidence  that:  (1)  The  content 

of  both  reducing  and  total  sugars  increases  in  hardened  plants.  This 
increase  seems  to  be  greater  in  plants  hardened  by  exposure  to  low 
temperature  in  the  coldframe  than  in  plants  hardened  by  other 
methods.  The  increase  in  sugar  is  greater  in  hardened  cabbage  and 
lettuce  than  in  the  tomato,  though  there  is  no  direct  evidence  that 
the  absolute  quantity  of  sugars  present  in  the  plant  is  directly  re- 
lated to  its  cold-resistance.  Thus  some  of  the  tender  lettuce  samples 
have  more  sugar  than  certain  samples  of  hardy  cabbage.  Young 
lettuce  plants  contained  much  more  sugar  than  plants  approaching 
maturity  and  this  may  have  something  to  do  with  the  greater  cold- 
resistance  Chandler20  found  in  the  younger  leaves  of  lettuce,  where- 
as in  most  plants  the  young  leaves  were  somewhat  more  tender  to 
cold. 

(2)  In  lettuce,  cauliflower  and  cabbage  the  amount  of  total 
polysaccharides  is  usually  somewhat  less  in  hardened  than  in  non- 
hardened  plants,  which  decrease  may  be  attributed  to  the  reduction 
in  the  amount  of  starch.  In  the  tomato,  on  the  other  hand,  the  total 
polysaccharides  show  a large  increase,  apparently  due  mostly  to 
the  deposition  of  starch  in  large  quantities  in  both  stems  and  leaves 
of  plants  exposed  to  any  of  the  hardening  treatments.  Kraus  and 
Kraybill54  found  a similar  increase  of  starch  in  tomato  plants  in  a 
stunted  condition.  Hartwell43  found  a large  accumulation  of  starch 
in  plants,  especially  the  potato,  when  the  growth  was  checked  by  any 
limiting  factor. 

Here  is  an  interesting  distinction  between  the  chemical  changes 
in  a group  of  plants  susceptible  of  considerable  hardening  to  cold 
and  a plant  not  susceptible  of  much  hardening.  In  the  group  of 
plants  possessing  potential  hardiness,  exemplified  by  the  cabbage, 
any  hardening  treatment  causes  a considerable  increase  in  sugars 
and  a decrease  in  starch,  while  the  total  polysaccharide  figure  re- 
mains nearly  constant  (on  the  fresh  weight  basis)  because,  as  will 
be  shown  later,  of  an  increase  in  pentosans.  On  the  other  hand  in 
the  tomato,  lacking  potential  hardiness,  the  hardening  treatments 
caused  only  a slight  increase  in  the  sugars  and  an  enormous  increase 
in  polysaccharides  due  mostly  to  an  increased  starch  content. 

An  increased  sugar  content  in  the  hardened  plants  would  in- 
crease the  osmotic  concentration  of  the  cell  sap,  depress  the  freez- 
ing point  and  perhaps  serve  to  hold  a somewhat  larger  amount  of 
water  in  the  unfrozen  state  when  the  plant  is  exposed  to  low  tem- 
peratures. However,  the  importance  of  the  increased  content  of  sim- 


66 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


pie  sugars  in  cold  resistance  remains  undetermined,  nor  is  it  known 
to  what  extent  sugars  may  be  responsible  for  the  greater  water-retain- 
ing capacity  of  hardened  tissues.  It  appears  probable  that  an  in- 
creased sugar  content  in  hardened  plants  is  more  likely  one  of  the 
manifestations  of  the  condition  of  being  hardy  than  a direct  cause  of 
cold  resistance. 

NATURE  OF  WATER-RETAINING  POWER  IN  PLANTS. 

It  has  been  shown  by  experiments  with  the  dilatometer  that: 
(1)  the  amount  of  water  frozen  in  hardened  cabbage  plants  is  con- 
siderably less  than  in  tender  plants,  (2)  the  increase  in  the  amount 
of  water  frozen  as  the  temperature  is  lowered  becomes  less  and  less, 
probably  approaching  zero,  (3)  the  amount  of  water  freezing  at  a 
given  temperature  (-5°C.)  decreases  as  the  degree  of  hardening  in- 
creases. It  has  also  been  shown  that  hardened  plants  have  a lower 
transpiration  rate  and  that  hardened  tissues  dry  out  more  slowly 
than  tender  tissues.  The  greater  water-retaining  power  of  the  cells 
of  hardened  plants  must  therefore  be  accepted  as  a fact.  What 
factors  are  responsible  for  the  development  of  this  increased  water- 
retaining  power? 

Several  investigators  have  attached  great  significance  to  the 
osmotic  concentration  of  the  sap  as  determined  by  the  depression  of 
its  freezing  point.  Some  data  are  also  presented  in  this  paper  (Table 
2)  showing  that  in  hardened  plants  this  depression  is  greater  than 
in  non-hardened  plants.  However,  concentration  of  the  sap,  even 
if  entirely  due  to  substances  having  a low  eutectic  point,  would  not 
be  sufficient  to  account  for  the  amount  of  water  found  to  remain 
unfrozen  in  hardened  plants  (Table  3).  Moreover,  some  of  the 
sap  solutes  have  a high  eutectic  point,  for  Harvey42  found  numerous 
large  crystals  of  calcium  malophosphate  in  frozen  spots  on  leaves 
exposed  to  temperatures  not  low  enough  to  kill  the  whole  plant.  Some 
investigators  have  stated  that  the  increased  sugar  content  of  plants 
in  winter  was  at  least  to  some  extent  responsible  for  their  cold-re- 
sistance because  of  the  increased  concentration  thereby  imparted  to 
the  cell  sap.  The  highest  percentage  of  total  sugar  found  in  cab- 
bage in  these  experiments  (Table  12,  1.461  percent  in  Series  El, 
gathered  March  22,  1920,)  is  equal  to  only  1.68  percent  sugar  solu- 
tion in  the  plant  sap.  Considering  half  of  this  sugar  to  be  glucose 
and  half  sucrose,  we  have  a sugar  solution  equivalent  to  less  than 
0.075  molecular.  This  would  not  be  sufficient  to  affect  materially  the 
amount  of  water  frozen  in  the  plant  tissue  at  a point  several  degrees 


The  Hardening  Process  in  Vegetable  Plants. 


67 


below  0°C.,  one  gram-molecular  weight  of  a non-electrolyte  lowering 
the  freezing  point  1.86° C. 

It  has  been  further  pointed  out  (p.  35)  that  the  greater  depres- 
sion of  the  freezing  point  in  hardy  tissues  is  associated  with  certain 
changes  in  the  plant  cell  upon  hardening,  the  apparent  increase  in 
sap  concentration  being  simply  an  accompaniment,  rather  than  a 
cause  of  increased  hardiness.  It  therefore  is  necessary  to  introduce 
some  other  factor  to  explain  the  difference  in  amount  of  water  freez- 
ing in  hardy  and  in  tender  tissues.  It  has  been  indicated  that  the 
force  of  imbibition  may  be  a powerful  factor  in  withholding  water 
from  freezing.  This  force  varies  inversely  to  the  water  content,  but 
probably  increases  more  rapidly  than  the  rate  of  decrease  in  water 
content,  as  indicated  by  the  slow  rate  of  drying  leaves  and  of  col- 
loidal materials  after  they  have  been  dried  out  to  a certain  extent. 
It  is  a pretty  well  recognized  fact  that  tissues  with  lower  water  con- 
tent are  more  resistant  to  killing  by  cold. 

Plant  protoplasm  is  not  a compound  of  definite  chemical  com- 
position or  even  constant  physical  condition,  but  a colloidal  mixture 
of  the  emulsoid  type  varying  in  consistency  from  a hydrosol  to  that 
of  a hydrogel  and  containing  different  substances  which  may  be  pres- 
ent in  greater  or  less  amounts  at  different  times  and  in  different 
organs.  According  to  Seif riz, 113  the  change  from  one  state  to  an- 
other is  dependent  upon,  or  coincident  with,  changes  in  physiologi- 
cal activity.  Thus,  in  the  eggs  of  Fucus,  he  found  a progressive  in- 
crease in  viscosity  with  decreasing  physiological  activity.  Straus- 
baugh,117  as  a result  of  recent  investigations  on  the  plum  in  Minne- 
sota, suggests  that  the  prolonged  dormancy  and  water-retaining  power 
which  he  found  in  hardy  varieties  is  due  to  a change  in  colloidal  prop- 
erties creating  an  increased  power  of  imbibition.  The  work  of  these  in- 
vestigators is  significant,  since  hardy  plants  are  usually  at  a low 
state  of  physiological  activity  at  the  time  of  their  greatest  cold  re- 
sistance. 

Water  of  imbibition  may  be  held  by  molecular  capillarity  or  in 
the  absorbed  condition  by  the  hydrophilous  colloids  of  the  plant  cells. 
Such  water  is  not  readily  available  for  freezing,  in  other  words,  the 
force  with  which  it  is  held  must  be  overcome  by  a considerable  force 
of  crystallization  before  it  can  be  drawn  from  the  cell  and  frozen. 
Me  Cool  and  Millar80  have  suggested  the  classification  of  plant  mois- 
ture as  “free’  or  easily  freezable  and  “unfree”  or  not  easily  freez- 
able, somewhat  as  Bouyouces  has  classified  soil  water.  Such  a classi- 
fication necessitates  setting  an  arbitrary  temperature  of  freezing,  the 


68 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


relative  amounts  of  free  and  unfree  water  varying  with  the  tempera- 
ture at  which  freezing  takes*  place.  Yet  it  is  convenient  for  our  pres- 
ent purpose  to  refer  to  free  and  unfree  water  in  the  sense  that  the 
latter,  for  one  reason  or  another,  remains  unfrozen  at  a given  tem- 
perature. 

It  seems  from  the  work  of  Bouyoucos10  and  of  McCool  and  Mil- 
lar80 that  the  unfree  water  is  held  to  a very  large  extent  in  the  ad- 
sorbed condition  by  protoplasmic  colloids.  The  water-retaining  power 
of  colloids  and  the  quantity  of  certain  colloidal  materials  in  the  cell 
are  thereby  suggested  as  an  explanation  of  increased  water-retaining 
power  and  cold-resistance  in  plants.  * 

Relation  of  pentosan  content  to  cellular  water-retaining  power. 
— Spoehr ’s  work116  on  cacti  suggests  that  pentosans  may  be  the  spe- 
cific substances  which  increase  the  water-retaining  power  in  hardened 
plants.  He  found  that  the  pentosan  content  of  Opuntia  increased 
considerably  under  xerophytic  conditions  and  suggested  that  the 
large  water-retaining  power  of  the  pentosans  is  largely  responsible 
for  their  well-known  ability  to  survive  under  such  circumstances. 
The  work  of  Livingston138  and  others  has  shown  that  the  osmotic 
pressure  in  cacti  and  other  desert  plants  is  no  greater  than  in  many 
mesophytes,  hence  this  factor  probably  plays  only  a small  part  in 
the  water-holding  power  of  most  xerophytic  plants. 

Spoehr  found  by  analysis  of  desert  plants  that  in  cells  under- 
going water  depletion,  other  polysaccharides  were  changed  to  pento- 
sans, of  which  the  plant  mucilages  are  largely  composed.  Thus,  4 ‘un- 
due loss  of  water  caused  a change  in  the  cell  whereby  the  amount  of 
water  it  may  hold  is  greatly  increased.  ” Mac  Dougal86  considers  a 
change  of  this  sort  to  be  the  basis  of  xerophytism.  Water  of  imbibi- 
tion was  found  by  Spoehr  to  be  closely  related  to  the  presence  of  the 
pentose  polysaccharides.  Pentosan  formation  increased  decided^ 
with  low  and  decreased  with  high  water  content.  From  April  till 
June,  while  the  weather  was  very  dry,  pentosans  made  up  from  9 
to  12  percent  of  the  dry  weight.  During  the  rainy  weather  of  July, 
the  pentosan  content  fell  to  4.39  percent  of  the  dry  weight,  increas- 
ing to  12.5  percent  again  in  the  dry  cool  weather  of  the  fall  and 
falling  to  4.37  percent  when  the  winter  rains  set  in. 

Not  all  cacti  possess  a large  pentosan  content.  Spoehr  gives 
analysis  of  two  species  of  Opuntia  growing  at  Tucson,  as  follows: 

Fresh  weight  basis 

% water  % total  % total  % total  % 


sugars  polysach.  pentose  pentosan 

0.  Versicolor  82.15  1.97  1.50  0.36  0.230 

O.  Phaeacantha  78.70  3.53  3.22  1.64  1.550 


The  Hardening  Process  in  Vegetable  Plants. 


69 


In  view  of  this  difference  in  composition,  it  may  be  significant 
that  0.  Pha^acantlia  is  listed  in  Bailey’s  Encj^clopedia  of  Horticul- 
ture as  a hardy  variety  and  is  reported  by  Shreve  to  grow  in  the 
mountains  about  Tucson  to  an  altitude  of  7500  feet. 

A striking  property  of  the  pentosans  is  their  power  of  swelling 
and  taking  up  an  enormous  amount  of  water,  which  the  hexose  poly- 
saccharides do  not  do  to  nearly  so  marked  a degree.  The  occurrence 
of  pectins  in  the  middle  lamella  of  the  cell  walls  is  well  known. 
Spoehr  believes  that  they  are  also  distributed  through  the  protoplasm 
and  are  used  for  a variety  of  purposes.  The  plant  nucleo-protein 
has  been  found  by  Levine  and  Jacobs136  to  contain  the  pentose  group 
as  part  of  the  nucleic  acid  radical.  Tollens137  showed  that  pentosans 
were  widely  distributed  in  plants  and  were  limited  to  no  special 
tissue,  but  abundant  in  roots,  stems,  leaves  and  seeds.  He  found 
further  that  pentosans  showed  all  possible  variations  as  to  solubility 
in  water.  Swartz119  obtained  a water-soluble  pentosan  from  Dulce. 
In  the  crude  form,  this  was  very  hygroscopic,  but  this  property  was 
lost  after  several  purifications.  She  found  that  the  hemi-celluloses  of 
ten  species  of  marine  algae  were  chiefly  pentosans  and  galactans  and 
concluded  that  pentosans  and  hexosans  very  commonly  occur  to- 
gether, not  only  intimately  associated,  but  chemically  combined. 
Mac  Dougal81  goes  so  far  as  to  state  that  the  “ plant  protoplasm  con- 
sists of  a comparatively  inert  base  of  pentosans — in  colloidal  com- 
bination with  proteins,  amino  acids,  lipins,  and  salts.” 

As  to  the  origin  of  pentosans,  Spoehr116  shows  that  pentoses  can 
be  formed  from  the  hexosans  as  the  first  product  of  oxidation.  This 
view  is  corroborated  by  the  work  of  Ravena  and  Cereser,102  who 
found  no  marked  variation  in  pentosan  content  during  the  period  of 
photosynthetic  activity,  but  when  the  carbohydrate  food  consisted 
entirely  of  dextrose,  the  amount  of  pentosans  increased  greatly,  es- 
pecially in  light.  The  probability  of  pentosan  formation  from  the 
hexosans  is  indicated  also  by  the  increased  pentosan  content  in  the 
presence  of  high  total  sugar  and  diminishing  starch,  as  shown  later 
in  this  paper. 

Davis,  Daish  and  Sawyer24  found  no  diurnal  variation  in  the 
pentosan  content  of  plants.  However,  they  found  that  the  amount 
of  pentosan  in  the  leaf  of  the  Mangold  ( Beta  vulgaris)  increased 
from  August  to  October. 

Hornby48  found  that  the  pectin  content  varied  in  different  parts 
of  the  same  plant.  More  pectin  was  found  in  the  epidermal  tissue 
than  in  the  cortex.  Exposure  to  light,  and  mechanical  injury  to 


70 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


tissues,  were. found  to  result  in  increased  pectin  content  in  the  ex- 
posed or  injured  part.  Hornby  suggested  that  pectin  might  have  a 
protective  effect  on  plants,  especially  against  insect  attacks. 

Hooker47  has  shown  that  the  hardier  parts  of  apple  shoots,  the 
bases,  have  a greater  water-retaining  power  than  the  tips,  which  are 
less  cold-resistant.  He  placed  portions  of  the  air-dried  ground  ma- 
terial in  desiccators  containing  sulfuric  acid,  the  concentration  of 
which  ranged  from  100  to  36.69  percent.  The  air-dry  material  lost 
moisture  in  the  desiccators  containing  the  higher  concentration  of 
acid  and  this  loss  was  greater  in  the  tender  material.  But  over  the 
lowest  concentration  of  acid  used,  water  was  taken  up,  the  gain  in 
weight  being  greater  in  the  hardy  material.  This  experiment  indi- 
cates that  hardy  apple  twigs  contain  a larger  amount  of  some  hygro- 
scopic material.  Hooker  attributed  the  greater  water-retaining  power 
of  the  hardy  tissue  to  the  larger  percentage  of  total  pentosan  found 
therein. 

Pentosan  content  in  the  hardening  process  in  vegetable  plants. 

— In  this  work,  a study  was  made  of  the  pentosan  content  in  an 
effort  to  throw  light  on  the  nature  of  the  increased  water-retaining 
power  of  hardened  plants.  For  the  pentosan  determinations,  sam- 
ples were  taken  from  plants  grown  under  the  various  hardening 
treatments  previously  described.  Also  a series  of  analysis  were 
made  on  plant  material  gathered  from  the  field  at  intervals  during 
the  fall  of  1920. 

Method  of  Pentosan  Analysis. — The  method  of  analysis  was 
that  employed  by  Spoehr.116  A two  gram  sample  of  the  oven-drv 
material  was  liydrolized  by  boiling  for  three  hours  with  eight  cc. 
concentrated  HOI  in  150  cc.  water.  After  cooling,  the  entire  con- 
tents of  the  flask  containing  the  products  of  hydrolysis  were  trans- 
ferred to  a 400  cc.  baker,  neutralized  with  NaOH,  a uniform  amount 
of  a suspension  of  yeast  was  added  and  the  beakers  placed  in  an 
oven  at  35-40 °C.  over  night.  The  hexose  sugars  were  fermented  off, 
Reaving  the  non-fermentable  pentose  sugars  in  the  solution.  After 
fermentation  the  material  was  filtered  and  washed,  the  filtrate  con- 
taining the  pentose  sugars  was  boiled  ten  minutes  to  drive  off  the 
alcohol,  then  prepared  for  analysis  in  the  same  way  as  described  for 
sugar  determinations.  The  result  obtained  was  calculated  from 
Munson  and  Walker’s  tables,  multiplying  the  glucose  value  by  0.85, 
since  Spoehr  found  that  the  reducing  value  of  the  pentose  sugars 
held  that  relation  to  glucose.  The  results  on  total  pentosan  content 
are  given  in  Table  13. 


The  Hardening  Process  in  Vegetable  Plants. 


71 


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72 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Plants  not  hardened  by  any  special  treatment  are  low  in  total 
pentosans  and  hardened  plants  have  a much  larger  amount,  in  some 
cases  in  cabbage  an  increase  of  about  200  percent.  Plants  given 
intermediate  hardening  treatments  have  a medium  amount  of  pen- 
tosans. The  increased  pentosan  content  of  the  hardened  plants  is 
most  striking  if  we  consider  the  results  on  the  fresh  weight  basis. 
This  probably  is  the  most  suitable  criterion  to  use  in  a study  of  the 
reactions  which  concern  the  living  plant,  especially  since  Parker  has 
shown  that  the  force  with  which  water  is  held  by  finely  divided 
materials  depends  largely  on  the  moisture  content. 

It  may  seem  that  the  absolute  amounts  of  pentosans,  even  in 
the  hardened  plants,  are  too  small  to  influence  very  markedly  the 
force  with  which  the  cells  may  retain  water  under  conditions  of 
stress.  However,  it  should  be  borne  in  mind  that  in  nature  the  pen- 
tose molecule  probably  exists  in  combination  with  four  molecules  of 
galactose  or  other  hexose  sugar.  Hence  the  amount  of  pentosans  in 
the  plant  is  much  greater  than  the  analyses  indicate. 

Pentosan  content  of  garden  plants. — Samples  of  leaves  were 
gathered  at  intervals  during  the  fall  from  cabbage,  kale  and  celery 
plants  growing  in  the  open  field.  The  seed  had  been  sowed  in  July 
and  the  plants  made  considerable  growth  before  the  first  light  frost 
came  on  October  1.  The  month  of  October  was  mild,  and  the  plants 
remained  alive  until  heavy  freezes  the  last  of  November.  Exposed 
to  steadily  declining  seasonal  temperatures,  these  plants  may  be  con- 
sidered to  have  undergone  a kind  of  hardening  treatment,  for  they 
were  able  to  withstand  light  frost  in  October  and  heavy  frost  the 
early  part  of  November.  The  results  of  the  total  pentosan  deter- 
minations are  given  in  Table  14. 

Table  14. — Total  Pentosan  Content  of  Garden  Plants  in  Autumn. 


Kale  Cabbage  Celery 


Date  sample 
collected 

% of 
fresh 
wt.  | 

I % of 
i dry 
wt.  I 

.%  fresh 
wt. 

% dry 
wt. 

%of 

fresh 

wt. 

% of 
dry 
wt. 

Sept.  15  

0.289 

4.06 

Oct.  7 

0.511 

3.93  ' 

0.580 

4.36  ! 

0.567 

4.42 

Oct.  20  

0.528  | 

4.89 

0.545 

4.73 

0.801  ' 

4.26 

Nov.  3 

0.537  i 

3.93 

0.621 

4.36  i 

0.793 

4.44 

Nov.  10  

Nov.  18  

0.722  ! 
1.064  j 

4.95 

6.48 

0.782 

! 5.31  ! 

1.029  | 

5.58 

Table  14  shows  that  the  total  pentosan  content  of  these  plants 
becomes  high  when  thej^  are  exposed  to  cool  weather  during  the  late 


'The  Hardening  Process  in  Vegetable  Plants. 


73 


fall.  The  pentosan  content  on  the  fresh  weight  basis  increases  fairly 
regularly  up  to  date  of  last  sampling. 

Pentosan  content  in  plants  watered  with  salt  solutions. — An 

experiment  wherein  the  hardiness  of  cabbage  plants  was  consider- 
ably increased  by  watering  them  with  M/10  salt  solutions  has  been 
described.  Plants  hardened  in  this  way  were  shown  to  have  greater 
water-retaining  power  than  unhardened  plants.  Samples  from  the 
salt  treatment  plots  were  analyzed  for  total  pentosan  content.  The 
results  are  given  in  Table  15. 

Table  15. — Pentosan  Content  in  Cabbage  Plants  Hardened  by  .Salt  Solu- 
tions. 


Treatment  of  plants 

Percent  total  pentosans 

On  fresh  weight  basis 

on  dry  wt.  basis 

Compost  soil,  tap  water  

0.290 

4.25 

Compost  soil,  NaNO 

0.471 

5.05 

Compost  soil,  KC1  

0.451 

4.24 

Compost  soil,  NaCl  

0.483 

5.05 

Sand,  tap  water  

0.220 

3.45 

Sand,  NaCl  

0.288 

4.25 

The  total  pentosan  content  of  the  plants  whose  growth  was 
checked  by  the  application  of  the  salt  solutions  and  which  were 
hardier  to  cold,  show  somewhat  greater  amounts  of  total  pentosans 
on  the  dry  weight  basis  and  a considerable  increase  on  the  fresh 
weight  basis,  as  compared  to  plants  making  a normal  growth  with 
tap  water. 

It  appears  from  Table  15,  that  the  pentosan  content  of  the  plants 
grown  in  sand  is  considerably  lower  than  for  plants  grown  in  com- 
post soil  and  receiving  corresponding  treatments.  The  plants  grown 
in  sand  and  receiving  tap  water  were  somewhat  tenderer  to  cold  than 
those  grown  in  compost  and  likewise  given  tap  water.  Plants  grown 
in  sand  and  watered  with  M/10  NaCl  show  only  a slight  increase  in 
pentosan  content,  as  compared  to  plants  grown  in  compost  soil,  like- 
wise watered  with  M/10  NaCl.  Here  again,  pentosan  content  shows 
a close  correlation  with  the  hardiness  of  the  plants,  as  determined  by 
freezing  experiments. 

Rate  of  increase  in  pentosan  content. — The  three  groups  of  ex- 
periments just  described  having  indicated  a larger  amount  of  pen- 
tosans in  plants  hardened  in  different  ways,  it  was  deemed  desirable 
to  determine  their  rate  of  development  during  the  hardening  process. 
Lots  of  potted  cabbage  plants  were  removed  from  the  warm  green- 


74 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


house  at  intervals  during  March,  and  placed  in  an  open  coldframe. 
On  March  19,  samples  were  taken  for  analysis  from  all  the  lots  which 
had  been  exposed  to  the  hardening  process  for  periods  ranging  from 
3 to  20  days,  as  well  as  from  some  of  the  original  lot  which  had  been 
kept  in  the  greenhouse  under  favorable  growing  conditions.  The 
total  pentosan  content  of  the  plants  hardened  for  varying  lengths  of 
time  is  given  in  Table  16. 


Table  16. — Rate  of  Increase  of  the  Total  Pentosan  Content  in  Cabbage 

Plants. 


Percent  pentosan 

Treatment 

On  fresh  weight 

On  dry  weight 

basis 

basis 

Greenhouse  plants,  not  hardened  1 

0.260 

2.97 

Hardened  in  frame  3 days  

0.374 

3.56 

Hardened  in  frame  5 days  

0.442 

3.86 

Hardened  in  frame  10  days  

0.750 

5.00 

Hardened  in  frame  20  days  

0.776 

5.84 

Pays  duration  of  Exposure  in  Coldfranc. 


Fig.  8. — Rate  of  increase  in  total  pentosan  content  of  cabbage  leaves  during  the  harden- 
ing process. 


The  results  of  Table  16  are  shown  graphically  in  Figure  8.  It 
appears  that  the  increase  in  pentosan  content  proceeds  quite  rapidly 
and  at  a fairly  uniform  rate  for  ten  days.  After  the  first  ten  days 
of  exposure  in  the  coldframe  the  pentosan  content  increased  only 


The  Hardening  Process  in  Vegetable  Plants. 


75 


slightly  in  this  experiment.  Other  experiments  have  shown  that  the 
cabbage  plant  acquires  nearly  its  maximum  degree  of  hardening  in 
this  time.  The  dry  matter  content  likewise  increases  rapidly  the  first 
few  days  of  the  hardening  process,  and  more  slowly  thereafter. 

In  the  dilatometer  experiments,  it  was  found  that  the  amount  of 
water  frozen  at  -5°C.  decreased  with  the  duration  of  the  hardening 
treatment  in  approximately  the  same  order  as  the  pentosan  content 
is  shown  to  have  increased  here.  This  seems  to  indicate  a close  relation- 
ship of  pentosan  content  to  water-retaining  power  and  to  cold  re- 
sistance. The  plants  used  in  the  dilatometer  experiments  were  of 
the  same  lots  as  those  from  which  the  pentosan  analyses  were  made. 


Table  17. — Relation  of  Hot-Water- Soluble  Pectins  to  Total  Pentosan 
Content  in  the  Hardening  Process. 


Percent  pentosan  on  fresh  weight  basis 

Treatment 

Date 

sample 

taken 

Total 

Hot-water 

soluble 

Insoluble 
(by  differ- 
ence) 

Cabbage 

Wet-grown  green- 
house plants 

3/12/20 

0.215 

0.075 

0.140 

Dry-grown  green- 
house plants 

3/12/20 

0.423 

0.292 

0.131 

Greenhouse  plants 
not  hardened 

3/22/20 

0.207 

0.091 

0.116 

Hardened  in  cold- 
frame  2 weeks 

3/22/20 

0.530 

0.408 

0.124 

Hardened  in  cold- 
frame  3 weeks 

3/16/21 

0.776 

0.550 

0.226 

Tomato 

Wet-grown  in 
greenhouse 

5/3/20 

0.693 

0.070 

0.623 

Dry-grown  in 
greenhouse 

5/3/20 

0.720 

0.071 

0.649 

Greenhouse  plants 
not  hardened 

5/3/20 

0.384 

0.051 

0.333 

Hardened  in  cold- 
frame  2 weeks 

5/3/20 

0.682 

0.071 

0.611 

Sweet  Potato 

Garden  plant 

10/7/20 

0.477 

0.127 

0.350 

Kale 

Garden  plant 

10/7/20 

0.511 

0.223 

0.288 

Garden  plant 

11/18/20 

1.064 

0.418 

0.646 

Celery 

Garden  plant 

10/7/20 

0.567 

0.236 

0.331 

Garden  plant 

1 11/10/20 

0.793 

0.423 

0.370 

76 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Relation  of  hot-water-soluble  pentosans  to  the  hardening  pro- 
cess.— In  jelly-making  a hot  water  extract  of  fruits  is  used.  Ac- 
cording to  Goldthwaite33  a cold  water  extract  of  our  common  fruits 
contains  little  or  no  pectin.  The  total  pentosan  determinations  given 
in  the  four  preceding  tables  indicate  the  larger  content  of  pentosans 
in  hardened  plants,  but  in  the  total  pentosans  is  included  probably 
a more  or  less  considerable  amount  of  the  insoluble  hemi-celluloses 
of  the  cell  wall,  which  might  not  be  expected  to  function  to  any  great 
extent  as  water-retaining  material,  though  undoubtedly  a part  of 
the  power  of  imbibition  of  the  plant  cell  is  due  to  its  walls.  The 
experience  of  jelly  makers  indicates  that  the  hot  water  extract  of 
fruit  contained  the  most  of  the  jelly-forming  pectins.  It  was  thought, 
therefore,  that  a hot  water  extract  of  the  plant  material  would  yield 
approximately  that  fraction  of  the  total  pentosan  which  exists  in 
the  protoplasm  and  might  function  as  the  significant  water-retain- 
ing material. 

Accordingly,  analyses  were  made  from  some  of  the  samples, 
varying  the  procedure  from  that  described  for  the  total  pentosan 
determinations  as  follows:  The  weighed  sample  of  dry  material  was 
transferred  to  a beaker  with  150  cc.  of  distilled  water.  The  slight 
acidity  was  neutralized  by  adding  a bit  of  sodium  carbonate,  then 
the  material  was  boiled  for  five  minutes,  and  filtered  hot  through  a 
Gooch  crucible.  This  yielded  a clear  cherry-colored  filtrate,  con- 
taining all  the  hot-water-soluble  pentosans,  sugars,  and  other  soluble 
carbohydrates.  Hydrolysis,  fermentation,  clearing  and  analysis  were 
carried  out  with  this  filtrate  in  the  same  way  as  previously  described 
for  the  whole  sample  in  the  total  pentosan  determinations.  The  re- 
sults are  given  in  Table  17. 

In  cabbage  plants  exposed  to  hardening  treatment,  the  water- 
soluble  pentosans  increase  considerably  while  the  insoluble  (hemi- 
cellulose)  fraction  is  nearly  constant,  regardless  of  the  degree  of 
hardiness.  In  hardened  cabbage  plants  the  amount  of  soluble  pen- 
tosans is  relatively  large,  in  fact  the  increase  in  the  total  pentosan 
content  is  very  largely  due  to  the  increase  in  the  water-soluble  frac- 
tion. 

In  tomatoes,  on  the  other  hand,  the  hot  water  soluble  fraction 
is  very  small  and  does  not  increase  much  in  plants  subjected  to 
hardening  treatments.  The  relatively  large  amount  of  total  pen- 
tosans in  the  tomato,  therefore,  is  largely  insoluble,  probably  exist- 
ing mostly  as  hemi-cellulose  or  in  the  middle  lamella.  The  sweet 
potato  resembles  the  tomato,  in  that  it  has  a relatively  large  total 


The  Hardening  Process  in  Vegetable  Plants. 


77 


pentosan  content,  but  only  a small  soluble  fraction.  Tlie  sweet  po- 
tato, like  the  tomato,  is  very  tender  to  frost  and  is  not  susceptible  of 
much  increase  in  cold-resistance  upon  exposure  to  usual  conditions 
of  hardening'. 

The  water-soluble  fraction  of  the  total  pentosan  content  in  gar- 
den plants  of  kale  and  celery  is  shown  to  increase  considerably  as 
they  become  hardier  in  the  fall. 

These  differences  in  the  soluble  pentosan  content  may  give  us 
an  important  clue  to  the  reason  for  the  previously  shown  difference 
in  cold-resistance,  susceptibility  to  hardening  and  water-retaining 
power  in  the  two  groups  of  plants  represented  respectively  by  the 
cabbage  and  the  tomato. 

Factors  influencing  the  imbibitional  capacity  of  plant  colloids. 

— In  view  of  the  increase  in  hardened  plants  of  pentosans,  especially 
in  the  hot-water-soluble  fraction,  and  the  possibility  of  these  sub- 
stances being  at  least  partly  responsible  for  the  increased  water-re- 
taining capacity  of  such  plants,  factors  which  influence  the  water- 
retaining  power  of  these  and  other  hydrophilous  colloids  occurring 
in  plants  may  be  of  great  importance  in  relation  to  cold-resistance. 


Fig.  9. — Swelling  of  Agar  as  influenced  by  reaction  of  the  solution. 


78 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Acidity. — Fischer30  showed  that  the  power  of  imbibition  of  col- 
loids was  influenced  very  markedly  by  the  reaction  of  the  medium, 
as  demonstrated  by  his  experiments  in  which  slight  acidity  increased 
the  swelling  of  gelatin.  He  was  able  to  alleviate  oedema  of  the  eye 
and  other  animal  tissues  by  application  of  alkali  and  hypertonic 
sugar  solutions.  Fischer  regards  acidosis  as  one  of  the  most  im- 
portant causes  of  the  presence  of  abnormal  amount  of  water  in  cells. 
Dachnowski23  found  that  seeds  of  beans  and  corn  swelled  more  and 
retained  more  w7ater  in  N/800  acids  than  in  water,  but  the  amount 
of  water  absorbed  and  retained  was  not  proportional  to  the  concen- 
tration of  acid,  for  a maximum  was  attained  beyond  which  increased 
acidity  decreases  absorption.  The  addition  of  equi-molecular  solu- 
tions of  non-electrolytes,  such  as  glucose  and  sucrose,  did  not  increase 
the  amount  of  water  retained  by  seeds  in  Dachnowski ’s  experiments. 
The  amino-acid,  glycocoll,  was  a striking  exception  in  that  greatly 
increased  imbibition  by  seeds  took  place  in  the  presence  of  this  sub- 
stance. Upson  and  Calvin141  have  shown  that  the  mixture  of  vegetable 
proteins  which  comprises  the  gluten  of  wheat,  behave  in  the  same 
way  as  Fischer’s  animal  proteins.  They  obtained  maximum  absorp- 
tion of  water  in  0.01  N hydrochloric  acid  and  0.04  N acetic  acid,  with 
marked  depression  of  absorption  by  strong  acids  and  by  salts.  Mac 
Dougal  and  Spoehr87  found  a greater  swelling  of  agar  in  N/100  solu- 
tions of  the  amino-acids  glycocoll,  alanin,  and  phenylalanin,  than  in 
water.  The  same  workers  have  shown  that  the  imbibition  of  protei- 
naceous colloids,  such  as  gelatin,  could  be  increased  considerably  by 
dilute  acids,  whereas  colloids  such  as  agar,  having  a pentosan  base, 
swelled  less  in  N/100  HC1  than  in  distilled  water. 

However,  brief  series  of  tests  made  by  the  writer  on  the  swelling 
of  agar  as  influenced  by  the  reaction,  indicate  that  the  greatest  swell- 
ing of  this  material  occurs  in  about  N/5000  HC1.  Presumably^  it 
would  require  a much  greater  concentration  of  the  plant  acids  to 
bring  about  the  same  degree  of  swelling  as  such  a dilute  HC1  solution 
Alkalinity,  excess  acidity,  and  the  presence  of  salts  depressed  the 
imbibitional  capacity  of  agar  very  markedly.  The  results  of  a du- 
plicate series  of  tests  performed  with  shredded  agar  are  presented 
graphically  in  figure  9. 

The  results  obtained  by  Mac  Dougal82  indicate  that  a mixture  of 
agar  and  gelatin  would  exhibit  maximum  swelling  in  somewhat 
stronger  acid  than  would  agar  alone.  Since  colloids  of  the  pentosan 
type  probably  occur  in  plants  in  intimate  association  with  proteina- 
ceous colloids,  it  is  reasonable  to  suppose  that  the  greatest  power  of 


The  Hardening  Process  in  Vegetable  Plants. 


79 


imbibition  would  be  exhibited  by  plant  cells  in  the  presence  of  slightly 
increased  acidity.  Mac  Dougal  and  Spoehr88  suggest  that  the  in- 
creased acidity  found  in  succulent  plants  may  be  a characteristic  of 
a metabolic  complex  favorable  to  pentosan  formation  and  to  the 
development  of  succulence  (a  high  degree  of  water-holding  power). 

In  connection  with  the  increased  water-holding  power  of  some 
colloids,  associated  with  slightly  increased  acidity  and  especially  some 
of  the  amino  acids,  it  is  interesting  to  note  that  Harvey42  found  a 
marked  increase  in  amino-nitrogen  in  hardened  cabbage.  May  it  not 
be  possible  that  in  developing  hardiness,  plants  form  some  specific 
amino  acid  which  would  increase  the  water-retaining  power  of  the 
cells  ? 

Somewhat  greater  titratable  acidity  has  been  found  in  hardened 
cabbage,  as  shown  in  Table  18.  Determinations  of  the  hydrogen-ion 


Table  18. — Titratable  Acidity  in  Hardened  and  Tender  Plants. 
(cc.  N/10  NaOH  per  one  gram  dry  material  in  100  cc.  water) 


Treatment 

Cabbage 

Lettuce 

(a) 

Tomato 

(b) 

Greenhouse  plants,  tender  . 

. . . 1.60 

1.86 

0.96 

1.74 

Coldframe  plants,  hardy  . . . 
In  coldframe,  2 weeks  

. . . 2.06 
. . . 1.68 

3.06 

1.74 

Grown  dry  in  greenhouse 

(hardy)  

Grown  wet  in  greenhouse 

. . . 1.30 

— t — 

0.96 

1.00 

(tender)  

. . . 0.82 

— 

0.66 

1.44 

concentration  were  also  made  on  a few  samples,  but  little  variation 
could  be  detected  by  the  Gillaspie  method. 

It  seems  that  there  is  a slight  increase  in  acidity  in  plants  as 
a result  of  the  hardening  process.  This  change  may  take  place  only 
in  plants  possessing  potential  hardiness  such  as  cabbage  and  lettuce 
since  the  data  in  Table  18  indicate  no  correlation  between  acidity  and 
hardening  treatments  in  the  tomato.  Increased  acidity  might  also 
influence  the  water-retaining  power  of  plant  cells  to  such  an  extent 
as  to  account  at  least  partly  for  the  cold-resistance  of  hardened  plants, 
aside  from  any  increase  in  the  amount  of  hydrophilous  cell  colloids. 
However,  too  few  data  are  available  to  draw  a definite  conclusion  on 
this  point. 

Salts  and  sugars. — It  has  been  shown  by  Fischer,  Dachnowski, 
Mac  Dougal  and  his  co-workers  that  the  addition  of  salts  to  a solution 
greatly  decreases  the  imbibitional  capacity  of  gelatin,  seeds  and 


80 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


agar.  However,  Free29  found  that  gelatin  swells  a little  more  in  0.5 
percent  solutions  of  dextrose  and  glucose  than  in  water,  while  a dis- 
tinct decrease  of  swelling  occurred  in  solutions  of  25  percent  or  over. 
Agar  was  found  to  swell  a little  more  in  two-percent  sucrose  than 
in  distilled  water,  whereas  dextrose  had  little  effect,  except  that  it 
depressed  swelling  in  concentrated  solutions.  That  dilute  sugar  so- 
lutions do  not  decrease  the  imbibitional  capacity  of  such  hydrophilous 
colloids  as  gelatin  and  agar  is  important,  since  a greater  sugar  content 
is  found  in  hardened  plants.  According  to  Goldthwaite33  pectin  and 
acid  are  prerequisites  for  j edification  of  fruit-juice,  while  sugar  is  a 
necessary  accessory.  She  was  able  to  make  an  excellent  artificial 
jelly  with  one  percent  pectin,  0.5  percent  tartaric  acid,  and  three- 
fourths  volume  of  cane  sugar.  Furthermore,  in  her  experiments,  it 
was  shown  that  increasing  the  proportion  of  sugar  gave  an  increased 
volume  of  jelly.  The  work  of  these  investigators  suggests  the  pos- 
sibility of  increased  acidity  and  sugar  content  playing  an  important 
part  in  determining  the  state  of  the  colloidal  protoplasm. 

Perhaps  sudden  or  extreme  changes  in  some  of  these  factors, 
which  influence  imbibitional  capacity,  might  exert  an  important  in- 
fluence on  the  water-retaining  power  and  cold  resistance  of  plant 
tissue.  However,  the  capacity  of  plant  organs  to  take  up  or  imbibe 
large  amounts  of  water  must  not  necessarily  be  taken  as  an  index  of 
their  power  to  retain  water  when  exposed  to  conditions  favoring  un- 
due water  loss,  such  as  freezing  or  drying. 

SUMMARY. 

The  work  of  previous  investigators  indicates  that  water-loss  from 
the  cells,  by  the  formation  of  ice  crystals  in  the  intercellular  spaces, 
is  most  generally  the  limiting  factor  in  the  killing  of  plant  tissue  by 
cold. 

Any  treatment  materially  checking  the  growth  of  plants  increases 
cold-resistance.  In  cabbage  and  related  plants,  hardiness  increases 
in  proportion  as  growth  is  checked.  In  tomato  and  other  tender  spe- 
cies, the  checking  treatments  resulted  in  relatively  slight  increase  to 
cold-resistance.  The  various  means  of  hardening  plants  in  these 
experiments  have  resulted  in  about  the  same  type  of  changes  within 
the  plant. 

Cabbage  plants  hardened  by  various  treatments  contain  a larger 
amount  of  “ unfree,  ” or  not  easily  frozen  water,  as  measured  by 
the  dilatometer.  The  increment  in  unfree  water  corresponds  to  the 


The  Hardening  Process  in  Vegetable  Plants. 


81 


extent  to  which  growth  is  checked,  both  of  these  paralleling  the  de- 
gree of  cold  resistance. 

The  amount  of  water  frozen  at  different  temperatures  in  leaves 
of  varying  hardiness  was  measured.  The  percentage  of  moisture 
frozen  in  hardened  cabbage  leaves  at  -3°C.  and  at  -4°C.  is  about 
two-thirds  of  that  frozen  in  tender  cabbage  leaves  at  the  same  tem- 
perature. The  actual  amount  of  water  remaining  unfrozen  at  a 
given  temperature  is  greater  in  hardened  than  in  tender  leaves,  al- 
though their  total  moisture  content  is  less. 

The  percentage  of  total  moisture  frozen  in  leaves  increases  for 
each  successive  degree  of  temperature  lowering,  but  the  increase  be- 
comes rapidly  smaller  and  smaller.  The  amount  of  water  remaining 
unfrozen  in  hardened  cabbage  leaves  is  approximately  a logarithmic 
function  of  the  temperature. 

Cabbage  plants  exposed  to  low  temperatures  in  a coldframe  for 
varying  periods  have  a progressively  smaller  amount  of  w^ater  freez- 
able at  -5°C.,  the  longer  they  are  exposed  to  hardening.  The  per- 
centage of  freezable  water  decreased  quite  rapidly  in  the  first  four 
days  after  removal  from  the  greenhouse,  more  slowly  from  four  to 
fourteen  days  and  very  slowly  thereafter*  The  rate  of  decrease  in 
percentage  of  freezable  water  coincides  with  the  observed  rate  of 
hardening.  In  other  words,  the  hardening  process  in  cabbage  plants 
was  accompanied  by  a proportional  increase  in  the  amount  of  water 
unfrozen  at  -5°C.  The  amount  of  water  frozen  at  -5°C.  is  somewhat 
less  in  plants  exposed  to  slight  wilting  at  midday. 

The  effects  of  watering  plants  with  M/10  salt  solutions  are  as- 
sociated with  a condition  of  mild  physiological  drought.  The  degree 
of  such  drought  is  proportional  to  the  concentration  of  the  soil  solu- 
tion, which  in  turn  is  influenced  by:  (a)  the  amount  of  water-soluble 
material  present  and  (b)  the  power  of  the  soil  to  hold  a large  part 
of  the  soil  moisture  unfree  in  the  pure  or  nearly  pure  state. 

Hardened  cabbage  plants  lose  less  moisture  by  transpiration  per 
unit  of  leaf  area  than  tender  plants,  under  the  same  conditions.  The 
amount  of  water  lost  by  transpiration  per  plant  for  a given  period  is 
much  less  in  hardened  cabbage  plants  than  in  non-hardened  plants 
of  the  same  age  because  of:  (a)  the  lower  rate  of  transpiration  and 
(b)  the  smaller  size  of  hardened  plants.  This  accounts  for  the  fact 
that  hardened  plants  can  be  transplanted  to  the  field  with  less  wilting. 

The  rate  of  water  loss  from  hardened  cabbage  leaves  dried  in 
an  oven  at  60°C.  is  much  less  than  that  from  leaves  of  tender  plants. 
In  tomato,  the  rate  of  drying  is  only  slightly  less  in  hardened  than 


82 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


in  non-kardened  plants.  Comparing  the  rate  of  water-loss  from  to- 
mato and  cabbage  leaves,  it  is  found  that  hardened  tomatoes  lose 
water  somewhat  faster  than  tender  cabbage  leaves. 

The  lesser  amount  of  water  lost  by  ice  formation,  the  lower  rate 
of  transpiration  and  the  slower  rate  of  water  loss  upon  drying  in 
hardened  cabbage  plants,  may  be  explained  by  the  hypothesis  that 
hardening  develops  an  increased  water-retaining  capacity.  The 
water-retaining  power  of  plant  cells  is  due  to:  (a)  Osmotic  concen- 
tration (b)  Imbibition,  and  may  be  increased  by  means  of  either  or 
both  of  these  factors. 

Osmotic  concentration  of  plant  cells  may  be  increased  by: 

(1)  Decreasing  the  total  water  content. 

(2)  Increasing  the  amount  of  osmotieally  active  sap  solutes. 

(3)  Decreasing  the  amount  of  free  water  or  conversely,  by  in- 
creasing the  amount  of  unfree  water  held  by  colloidal  adsorption. 

Osmotic  concentration  as  measured  by  the  lowering  of  the  freez- 
ing point  has  been  found  to  increase  on  hardening  plants,  varying 
inversely  with  the  water  content.  Both  reducing  and  non-reducing 
sugars  increase  with  hardening.  Sugars  are  found  to  increase  more 
in  cabbage  and  lettuce  than  in  tomato.  The  increased  sugar  is  not 
sufficient  to  account  for  much  difference  in  the  freezing  point  depres- 
sion or  in  the  amount  of  water  remaining  unfrozen  several  degrees 
below  the  freezing  point.  The  chief  factor  in  increasing  osmotic 
concentration  in  plants  is  considered  to  be  the  decrease  in  amount  of 
free  water,  hence  the  observed  increase  in  osmotic  concentration  would 
be  a secondary  result  of  the  hardening  process. 

The  power  of  imbibition  possessed  by  plant  cells  may  be  increased 

by: 

(1)  Decreasing  the  total  water  content  (or  increasing  the  per- 
cent of  dry  matter). 

(2)  Increasing  the  amount  of  hydrophilous  colloids  in  the  pro- 
toplasm. 

(3)  Increasing  the  water-retaining  power  of  such  colloids  by 
slight  increase  in  acidity,  etc. 

Decreased  water  content  accompanies  a condition  of  greater 
cold  resistance  in  plants.  During  the  hardening  process,  the  per- 
centage of  dry  matter  increases  rapidly  for  a few  days,  and  more 
slowly  thereafter.  The  total  pentosan  content  is  greater  in  hardened 
th&n  in  tender  plants,  regardless  of  the  kind  of  hardening  treatment. 
The  pentosan  content  of  cabbage  plants  exposed  to  low  temperatures 
in  an  open  coldframe  during  March  increases  rapidly  the  first  five 


The  Hardening  Process  in  Vegetable  Plants. 


83 


days  and  more  slowly  thereafter.  The  pentosan  content  of  cabbage, 
kale  and  celery  plants  growing  in  the  open  garden  increases  as  the 
weather  becomes  colder  during  the  fall.  In  cabbage,  kale  and  let- 
tuce plants  possessing  potential  hardiness,  the  fraction  of  the  pen- 
tosan content  soluble  in  hot  water  is  larger  than  in  tomato,  eggplant 
and  sweet  potato,  which  do  not  possess  potential  hardiness.  The 
hot  water-soluble  pentosan  content  is  thought  to  represent  more 
nearly  the  amount  of  pentosans  in  the  protoplasm  and  these  might 
function  more  specifically  as  water-retaining  material.  In  the  group 
of  plants  susceptible  of  considerable  hardening  to  cold  the  increase 
in  total  pentosan  content  upon  hardening  is  largely  an  increase  in 
the  hot  water-soluble  fraction,  while  in  the  tomato  the  hot  water- 
soluble  fraction  does  not  increase  upon  subjecting  the  plants  to  har- 
dening treatments. 

CONCLUSIONS. 

The  experimental  data  show  that  the  hardening  process  in  plants 
is  accompanied  by  a marked  increase  in  water  retaining  power,  and 
that  this  water  retaining  power  is  due  chiefly  to  the  imbibitional 
forces  of  the  cell.  The  amount  of  water  frozen  in  hardy  plants  is 
less  than  in  tender  plants  and  cells  of  hardy  plants  actually  retain  a 
larger  amount  of  unfrozen  water  than  those  of  tender  plants. 

It  is  believed  that  cold  resistance  in  plants  is  due  to  the  increased 
water-retaining  power  of  the  cells,  which  enables  them  upon  freezing 
to  retain  a larger  proportion  of  their  moisture  content  in  the  unfrozen 
condition. 

The  increased  water-retaining  power  of  hardened  plants  is  as- 
sociated with  the  following  changes:  (a)  decreased  moisture  con- 

tent, (b)  increased  amount  of  hydrophilous  colloids,  such  as  pen- 
tosans, (c)  increased  water-retaining  power  of  such  cell  colloids  be- 
cause of  a slight  increase  in  acidity  or  other  internal  changes,  (d) 
increased  amount  of  osmotically  active  substances  as  soluble  sugars. 
The  last  factor  probably  is  important  only  in  plants  hardened  by 
prolonged  exposure  to  cold ; the  first  three  factors  mentioned  may 
become  operative  in  a very  short  time,  when  the  activity  of  the  plant 
is  limited  by  any  factor.  Perhaps  the  same  changes  which  increase 
the  water-retaining  power  also  favor  greater  stability  of  the  proto- 
plasm. 

The  marked  parallelism  between  pentosan  content  and  hardiness 
indicates  a causal  relationship.  However,  pentosan  content  alone  is 
not  to  be  taken  as  an  absolute  index  of  cold  resistance,  since  several 


84 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


factors  may  affect  tlie  functioning  of  pentosans  as  water-retaining 
substances.  Salt  content,  acidity,  hydrogen-ion  concentration,  sugar, 
moisture,  protoplasmic  colloids  other  than  pentosans  and  perhaps, 
other  factors  constitute  a varying  complex  which  may  influence  water- 
retaining  power  and  hardiness. 

The  differential  reactions,  when  subjected  to  hardening  treat- 
ments, of  plants  possessing  potential  hardiness  as  the  cabbage  and 
of  plants  lacking  it  as  the  tomato,  indicate  that  the  fundamental 
difference  between  hardy  and  tender  species  lies  in  their  ability  to 
initiate  changes  whereby  the  stability  and  water-retaining  power  of 
the  protoplasm  and  consequently  hardiness  are  increased.  Hardy 
species  and  varieties  of  plants  possess  the  ability  to  initiate  such 
changes  to  a greater  or  less  great  degree,  while  tender  species  pos- 
sess it  to  a very  slight  degree  or  not  at  all. 

APPLICATIONS. 

\ 

In  view  of  the  connection  between  cell  water  retaining  power 
and  hardiness  which  has  been  found  and  the  correlation  between 
soluble  pentosan  content  and  hardiness,  it  seems  that  problems  deal- 
ing with  cold  resistance  of  vegetables,  cereals,  fruits  and  shrubs  may 
be  attacked  from  a new  angle. 

Furthermore,  the  association  of  water-retaining  power  of  cells 
with  their  content  of  a specific  material  or  group  of  materials,  such 
as  ipijlfoguis,  may  be  important  in  the  study  of  moisture  relations 
an(^^vfTter!4novement  in  plants.  Moreover  it  may  lead  to  a better 
under standing\of^the  cause  and  prevention  of  a group  of  physiologi- 
cal plant  diseases  usually  associated  with  excessive  water  loss,  such 
as  Tipburn  of  potato  and  lettuce,  and  Blossom  End  Rot  of  tomato. 
Selection  of  plants  for  high  soluble  pentosan  content  may  be  helpful 
to  the  breeder  of  cold-resistant,  drought-resistant,  or  disease-resistant 
varieties  of  crop-plants. 

The  changes  of  the  food  value  of  fruits  and  vegetables  subjected 
to  long  storage  may  be  significant,  since  it  seems  that  in  living  plant 
tissues,  exposed  to  water  deficit  or  to  cold  the  hexosan  carbohydrates 
are  converted  into  pentosans,  which  have  a much  lower  coefficient  of 
digestibility.  However,  the  use  of  such  vegetables  as  have  a high 
water-retaining  power  may  be  important  dietetically  in  the  alleviation 
of  certain  digestive  disorders. 


The  Hardening  Process  in  Vegetable  Plants. 


85 


ACKNOWLEDGMENTS 

To  Messrs.  V.  R.  Gardner,  H.  D.  Hooker,  Jr.,  and  F.  C.  Brad- 
ford of  the  Department  of  Horticulture  and  W.  J.  Robbins  of  the 
Department  of  Botany,  I am  indebted  for  suggestions,  constructive 
criticisms  and  some  of  the  references  of  the  literature.  The  work  on 
measurement  of  cells  and  on  the  swelling  of  agar  was  performed  in 
the  Botanical  Laboratories  under  the  direction  of  Dr.  Robbins. 

The  writer  wishes  to  acknowledge  gratefully  the  assistance,  sug- 
gestions, criticisms,  and  kindly  encouragement  so  generously  extended 
by  all  of  these  gentlemen,  without  which  this  work  could  never  have 
been  performed. 


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pflanzliche  Zelle.  Mitt.  Kaiser  Wilhelms  Inst.  Landw.  Bromberg,  3: 
pp.  93-115,  1910. 

111.  Schulz,  E.,  Uber  Preserwestoffe  in  immergrunen  Blattern,  Flora,  71: 
p.  223,  1888. 

112.  Schimper,  A.  F.  W.,  Plant  Geography  upon  a Physiological  Basis. 
Trans,  by  W.  R.  Fischer,  p.  25-41,  The  Clarendon  Press.  Oxford,  1903. 

113.  Seifriz,  William,  Viscosity  of  Protoplasm  as  Determined  by  Microdis- 
section. Bot.  Gaz.  70:  pp.  360-378,  1920. 

114.  Schutt,  F.  T.,  On  the  Relation  of  Moisture  Content  to  Hardiness  in 
Apple  Twigs,  Proc.  & Trans.  Royal  Soc.  Canada  11,  9:  Sec.  IV,  pp.  149- 
153,  1903. 

115.  Sinz,  E.,  Beziechungen  zwischen  Trocksubstanz  und  Winterfestighalt 
bei  verschiedenen  winter-weizen  Varietatur.  Jour.  Landw.  62,  pp.  301- 
335,  1914.  (Abs.  in  Exp.  Sta.  Record,  33:  235,  1915.) 

116.  Spoehr,  H.  A.,  Carbohydrate  Economy  of  Cacti.  Publication  287,  Car- 
negie Inst.  Washington,  1919. 

117.  Strassbaugh,  P.  D.,  Dormancy  and  Hardiness  in  the  Plum.  Bot.  Gaz. 
71:  pp.  337-357,  1921. 

118.  Storber,  J.  P.,  Comparative  Study  of  Winter  and  Summer  Leaves  of 
Various  Herbs.  Bot.  Gaz.  63:  pp.  89-111,  1917. 

119.  Swartz,  Mary  D.,  Nutrition  Investigations  on  the  Carbohydrates  of 
Lichens,  Algae,  and  Related  Substances,  Trans.  Conn.  Acad.  Arts  and 
Sci.  16:  pp.  247-382,  1911. 

120.  Tuttle,  G.  M.,  Induced  Changes  in  Reserve  Materials  in  Evergreen 
Herbaceous  Leaves,  Ann.  Bot.  33:  pp.  201-210,  1919. 

121.  Uphof,  J.  C.  Th.,  Cold  Resistance  in  Spineless  Cacti.  Ariz.  Agr.  Exp. 

Sta.  Bui.  70,  1916. 

122.  Vass,  A.  F..  Influence  of  Low  Temperature  on  Soil  Bacteria.  Cornell 
Univ.  Agr.  Exp.  Sta.  Memoir  Bui.  27,  1919. 

123.  Voightlander,  H.,  Unterkuhlung  und  Kaltetod  der  pflanzen,  Beitr,  Biol. 
Pflanzen,  Bd.  9,  Heft,  31. 

124.  Walster,  H.  L.,  Formative  Effect  of  High  and  Low  Temperature  upon 
Growth  of  Barley,  Bot.  Gaz.  69:  pp.  97-126,  1920. 

126.  Weaver,  J.  E.  and  Morgensen,  A.,  Relative  Transpiration  of  Coniferous 
and  Broad  Leaved  Trees  in  Autumn  and  Winter.  Bot.  Gaz.  68:  pp.  393- 

424,  1918. 

127.  Webber,  H.  J.  et  al.  Effect  of  Freezes  on  Citrus  in  California.  Calif. 
Agr.  Exp.  Sta.  Bui.  304,  1919. 

128.  West,  F.  L.,  and  Edlefsen,  N.  E.,  Freezing  of  Peach  Buds.  Utah  Agr. 
Exp.  Sta.  Bui.  151. 


90 


Missour  Agr.  Exp.  Sta.  Research  Bulletin  48. 


129.  Wiegand,  K.  M.,  Some  Studies  Regarding  the  Biology  of  Buds  in 
Winter.  Bot.  Gaz.,  41:  pp.  373-424,  1906. 

130.  , Occurrence  of  Ice  in  Plant  Tissue.  Plant  World  9:  p. 

25,  1906. 

131.  , The  Passage  of  Water  From  the  Plant  Cell  During  Freez- 

ing. Plant  World,  9:  pp.  107-118,  1906. 

132.  Wright,  R.  C.  and  Taylor,  G.  F.,  Freezing  Injury  to  Potatoes  when 
Undercooled..  U.  S.  Dept.  Agric.,  Dept.  Bui.  916,  1921. 

133.  Dixon,  H.  H.,  Transpiration  and  the  Ascent  of  Sap.  In  Prog.  Rei. 
Bot  3:  pp.  1-66,  1910. 

134.  Drabble,  E.  and  Drabble,  H.,  The  Osmotic  Strength  of  Cell  Sap  in 
Plants  growing  under  Different  Conditions.  New  Phytologist,  4:  pp. 
189-191,  1905. 

135.  Ewart,  A.  J.,  On  the  Power  of  Withstanding  Desiccation  in  Plants. 
Proc.  Liverpool  Biol.  Soc.  11:  pp.  151-159,  1897. 

136.  Levene  J.,  and  Jacobs,  W.,  Ueber  die  Pankreas-Pentose,  Ber.  d.  deut. 
Chem.  Gesell,  43:  3147-3150,  1910. 

137.  Tollens.  Untersuchungen  uber  Kohlenhydrate.  Landw.  Versuchs-Sta- 
tionen,  39:  p.  401,  1891.  (Quoted  by  Swartz,  see  Bib.  No.  119). 

138.  Livingston,  E.,  Role  of  Diffusion  and  Osmotic  Pressure  in  Plants. 
Pamphlet,  75p.,  Chicago,  1903. 

139.  Reinke,  Quoted  by  Pfeffer-Physiology  of  Plants,  Vol.  1,  p.  73. 

140.  Pfeffer,  W.,  Physiology  of  Plants,  Second  Edition.  English  Trans, 
by  A.  J.  Ewart,  Vol.  1,  p.  73-75,  Oxford,  1900. 

141.  Upson  F.  W.  and  Calvin,  J.  W.,  The  Colloidal  Swelling  of  Wheat 
Gluten  in  Relation  to  Milling  and  Baking.  Nebraska  Agr.  Exp.  Sta., 
Research  Bui.  8. 


The  Hardening  Process  in  Vegetable  Plants 


91 


Plate  1. — Effect  of  Exposure  in  Open  Frames  on  Coed  Resistance. 

A.  Coldframe  hardened  vs.  greenhouse  plants  frozen  at  -4°C.  for  2 '/,  hours. 

Nov.  17,  1919. 

Vi.  Cabbage  plants  after  freezing  at  -8°C.  for  2 yz  hours,  March  28,  1921. 

(1)  Hardened  in  coldframe  two  weeks.  Lower  leaves  broken  off 
for  samples. 

(2)  Non-hardened  greenhouse  plant. 


92 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Plate  2. — Effect  of  Variation  in  Soil  Moisture  on  Cold  Resistance  of 

Cabbage. 

A.  Plants  grown  in  greenhouse  with  varying  supply  of  water;  after  freez- 
ing at  -4°C.  for  2y2  hours.  Nov.  17,  1919.  (1)  Dry  grown  (2)  Medium 
dry  (3)  Wet  grown. 

B.  (1)  Medium-dry-grown  greenhouse  cabbage  plants. 

(2)  Medium  wet  grown  greenhouse  cabbage  plants  after  freezing  at 
-4°C.  for  30  minutes,  March  28,  1921. 

C.  After  freezing  at  -4°C.  for  30  minutes,  March  28,  1921. 

(1)  Watered  heavily  until  one  week  before  this  test,  thereafter 
wilted  slightly  for  five  days. 

(2)  Plant  from  same  batch  as  (1)  but  not  subjected  to  preliminary 
wilting. 


The  Hardening  Process  in  Vegetable  Plants. 


93 


Plate  3. — Effect  of  Varying  Soil  Moisture  on  Hardiness  of  Tomato. 

A.  Greenhouse  tomato  plants  after  freezing  at  -2°C.  for  2 hours.  Sept.  29, 

1919. 


II. 


(1)  Dry-grown 

Greenhouse  tomato  plants  after 
Sept.  22,  1921. 

(1)  Dry-grown 


(2)  Wet-grown, 
freezing  at  -2.25°C.  for 


2/4 


hours, 


(2)  Medium-drv-grown 


(3)  Wet-grown. 


94 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Plate  4 — .Effect  of  Watering  Plants  Grown  in  Sand  in  Greenhouse  With 

M/10  Salt  Solutions. 

A.  After  freezing  at  -6°C.  for  30  minutes. 

(1)  NaCl  (2)  KC1  (3)  NaNO, 

B.  After  freezing  at  -3°C.  for  30  minutes 

(1)  NaCl,  (2)  KC1.  (3)  NaNO, 


(4)  Tap  water 
(4)  Tap  water 


The  Hardening  Process  in  Vegetable  Plants. 


95 


Pi.ate  5. — Effect  of  Watering  Cabbage  Plants  Grown  in  Greenhouse  With 

M/10  Salt  Solutions, 

A.  Grown  in  compost  soil  and  watered  with: 

(1)  NaCl  (2)  KC1  (3)  NaNO,  (4)  Tap  water. 

After  freezing  at  -6°C.  for  30  minutes. 

II.  Grown  in  Compost  soil  plus  rotten  manure,  watered  with: 

(1)  NaCl  (2)  KC1  (3)  NaNO:f  (4)  Tap  water. 

After  freezing  at  -6°C.  for  30  minutes. 


96 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  48 


Plate  6. — Relative  Wilting  of  Hardened  and  Tender  Cabbagei  Plants. 

A.  Cabbage  plants  from  transpiration  experiment  No.  4 after  5 hours  of 
exposure  to  high  transpiration  conditions. 

(1)  Greenhouse  plant,  watered  with  M/10  NaCl,  hardy. 

(2)  Greenhouse  plant,  watered  sparingly  with  tap  water,  hardy. 

(3)  Hardened  in  coldframe  5 days,  hardy. 

(4)  Greenhouse  plant  watered  heavily,  tender. 

B.  Coldframe  hardened  cabbage  plants  C.  Greenhouse  non-hardened 

one  day  after  transplanting  to  field,  plants,  handled  other- 

March  27,  1918,  weather  fair,  warm,  wise  the  same  as  those 

dry.  in  B. 


The  Hardening  Process  in  Vegetable  Plants. 


97 


Plate  7. — Tender  Cabbage  Plant  From  Greenhouse  Frozen  at  -5°C.  Fob 
30  Minutes,  March  31,  1921. 

Droplets  of  water  exuding  from  stem  and  petioles  upon  thawing.  The 
leaves  were  covered  with  a film  of  smaller  droplets. 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 


AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  49 


EXPERIMENTS  IN  FIELD  PLOT 
TECHNIC  FOR  THE  PRELIMINARY 
DETERMINATION  OF  COMPARATIVE 
YIELDS  IN  THE  SMALL  GRAINS 

(Publication  authorized  December  2,  1921.) 


COLUMBIA,  MISSOURI 
DECEMBER,  1921 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY,  P.  E.  BURTON,  H.  J.  BLANTON, 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 
J.  C.  JONES,  Ph.  D.,  LL.D.,  PRESIDENT  OF  THE  UNIVERSITY 


AGRICULTURAL 


STATION  STAFF 
DECEMBER,  1921 
CHEMISTRY 


RURAL  LIFE 


O.  R.  Johnson,  A.  M. 
S.  D.  Gromer,  A.  M. 
E.  L.  Morgan,  A.  M. 


C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  A.  M. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  Sieveking,  B.  S.  in  Agr. 

AGRICULTURAL  ENGINEERING 

J.  C.  Wooley,  B .S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

E.  F.  Hopkins,  Ph.  D. 

DAIRY  HUSBANDRY 

A.  C.  Ragsdale.  B.  S.  in  i\gr. 

W.  W.  Swett,  A.  M. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride, 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  A.  M. 

O.  W.  Letson,  B.  S.  in  Agr. 

B.  M.  King,  B.  S.  in  Agr. 

A.  C.  Hill,  B.  S.  in  Agr. 

Miss  Bertha  C.  Hite,  A.  B.1 
Miss  Pearl  Drummond,  A.  A.1 


Ben  H.  Frame,  B.  S.  in  Agr. 


horticulture 

V.  R.  Gardner,  M.  S.  A. 

H.  D.  Hooker,  Jr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  M.  S. 

F.  C.  Bradford,  M.  S. 

H.  G.  Swartwout,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY 

H.  L.  Kempster,  B.  S. 

Earl  W.  Henderson 


SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M. 

W.  A.  Albrecht,  Ph.  D. 

F.  L-  Duley,  A.  M.3 
R.  R.  Hudelson,  A.  M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr. 
Richard  Bradfield,  A.  B. 

O.  B.  Price,  B.  S.  in  Agr. 

veterinary  science 

J.  W.  Connaway,  D.  V.  S.,  M.  D. 
L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 


OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Sercretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Jane  Frodsham,  Librarian 
E.  E.  Brown,  Business  Manager 


*In  service  of  U.  S.  Department  of  Agriculture,  Seed  Testing  Laboratory. 
2On  leave  of  absence. 


CONTENTS 


Page 

The  Problem  6 

Plan  and  Method  of  Investigation  9 

Terminology  9 

Procedure  11 

Work  of  1919  12 

Work  of  1920  16 

Work  of  1921  18 

Competition  as  a Source  of  Error  in  Preliminary  Tests  23 

Previous  Investigation  23 

Experimental  Results  25 

Illustrations  of  Effects  of  Competition  26 

Relation  of  Competition  to  Various  Characteristics  of  the  Com- 
peting Varieties  31 

Discussion  40 

Size  and  Replication  of  Plots  43 

Previous  Investigation  43 

Experimental  Results  44 

Size  of  Plots  44 

Replication  of  Plots  50 

Adjustment  of  Yields  by  Means  of  Check  Plots  54 

Previous  Investigation  54 

Experimental  Results  56 

Method  Used  in  Adjusting  Yields  58 

Relative  Variability  of  Actual  and  Adjusted  Yields  60 

Difference  in  Results  Obtained  by  Adjustment  with  Different  Check 

Varieties  63 

Value  and  Limitations  of  Adjusting  Yields  by  Means  of  Check  Plots  71 

Concluding  Remarks  73 

Summary  75 

Acknowledgment  77 

References  Cited  78 


TABLES 

Table 

Number  Table  Page 

1 Yields  of  Barley  Varieties  1919  13 

2 Yields  of  Oats  Varieties  1919  14 

3 Yields  of  Oats  Strains  1919  15 

4 Yields  of  Wheat  Varieties  1920  16 

5 Yields  of  Wheat  Varieties  1921  17 

6 Yields  of  Wheat  Varieties  and  Mixtures  1921  18 

7 Yields  of  Oats  Varieties  1921  21 

8 Yields  of  Oats  Strains  1921  22 

9 Relative  Yields  of  Two  Small  Grain  Varieties  When  Compared  in  Al- 

ternate Rows  and  in  Blocks  (Kiesselbach)  24 

10  Correlation  of  Competition  with  Various  Characteristics  in  Barley  Va- 

riety Test  1919  35 

11  Correlation  of  Competition  with  Various  Characteristics  in  Oats  Va- 

riety Test  1919  35 


4 

12 

13 

14. 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 


Tables 


Correlation  of  Competition  with  Various  Characteristics  in  Oats  Strain 

Test  1919  36 

Correlation  of  Competition  with  Various  Characteristics  in  Wheat  Va- 
riety Test  1920  . . 37 

Correlation  of  Competition  with  Various  Characteristics  in  Wheat  Va- 
riety Test  1921  37 

Correlation  of  Competition  with  Various  Characteristics  in  Wheat  Mix- 
ture Test  1921  38 

Correlation  of  Competition  with  Various  Characteristics  in  Oats  Va- 
riety Test  1921  39 

Summary  of  Effects  of  Competition  in  All  Tests  41 

Correlation  of  Yield  with  Dates  of  Heading  and  Maturity  in  Variety 

Tests  of  Barley,  Oats,  and  Wheat  42 

Yield  and  Variability  of  1-row,  3-row,  and  5-row  Check  Plots  in  Bar- 
ley Variety  Test  1919  45 

Yield  and  Variability  of  1-row,  3-row,  and  5-row  Check  Plots  in  Oats 

Variety  Test  1919  46 

Yield  and  Variability  of  1-row,  3-row,  and  5-row  Check  Plots  in  Oats 

Strain  Test  1919  47 

Yield  and  Variability  of  1-row,  3-row,  and  5-row  Check  Plots  in  Wheat 

Variety  Test  1920  47 

Yield  and  Variability  of  3-row  and  5-row  Check  Plots  in  Wheat  and 

Oats  Test  1921  48 

Yield  and  Variability  of  3-row  and  5-row  Test  Plots  in  All  Tests  50 

Relation  of  Plot  Variability  to  Size  of  Experiment  Field  in  Wheat  Va- 
riety Test  1920  51 

Relation  of  Plot  Variability  to  Size  of  Experiment  Field  in  Wheat  Va- 
riety Test  1921  52 

Relation  of  Plot  Variability  to  Size  of  Experiment  Field  in  Oats  Va- 
riety and  Strain  Tests  1921  52 

Soil  Heterogeneity  of  an  Experiment  Field  as  Determined  from  Yields 

of  Two  Check  Varieties  53 

Effect  on  Plot  Variability  of  Adjusting  Yields  by  Check  Plots  (Kies- 

selbach)  55 

Reduction  of  Variability  by  the  Use  of  Check  Plots  Equivalent  to  That 
Probably  Attainable  with  the  Same  Number  of  Plots  by  Replication  . 57 
Relative  Variability  of  Actual  and  Adjusted  Yields  in  Barley  Variety 

Test  1919  59 

Relative  Variability  of  Actual  and  Adjusted  Yields  in  Oats  Variety 

Test  1919  60 

Relative  Variability  of  Actual  and  Adjusted  Yields  in  Oats  Strain 

Test  1919  61 

Relative  Variability  of  Actual  and  Adjusted  Yields  in  Wheat  Variety 

Test  1920  62 

Relative  Variability  of  Actual  and  Adjusted  Yields  in  Wheat  Variety 

Test  1921  64 

Relative  Variability  of  Actual  and  Adjusted  Yields  in  Wheat  Mixture 

Test  1921  ....65 

Relative  Variability  of  Actual  and  Adjusted  Yields  in  Oats  Variety 

Test  1921  66 

Relative  Variability  of  Actual  and  Adjusted  Yields  in  Oats  Strain 

Test  1921  67 

Relative  Variability  of  Actual  and  Adjusted  Yields  of  Kherson  and  Red 
Rustproof  Oats  Each  in  120  Distributed  Plots,  in  Oats  Variety  and 

Strain  Tests  1921  68 

Summary  of  Relative  Variability  of  Actual  and  Adjusted  Yields  of 
Interior  Rows  in  All  Tests  1921  71 


EXPERIMENTS  IN  FIELD  PLOT  TECHNIC 
FOR  THE  PRELIMINARY  DETERMINA- 
TION OF  COMPARATIVE  YIELDS  IN 
THE  SMALL  GRAINS* 

L.  J.  Stadler 

During  recent  years  the  investigation  of  the  reliability  of  field 
experiments  has  become  an  important  phase  of  agronomic  research. 
Field  experiments  as  ordinarily  conducted  have  been  shown  to  be 
affected  by  many  gross  errors.  In  the  light  of  these  investigations  it 
has  become  apparent  that  the  results  of  many  of  the  older  experiments 
are  inconclusive  or  even  misleading.  Various  expedients  have  been 
suggested  for  counteracting  experimental  error.  Some  of  these  have 
been  quite  successful,  while  others  have  probably  done  more  harm 
than  good. 

The  pioneer  investigations  in  this  field  have  been  of  great 
value  in  directing  attention  to  the  important  sources  of  error  and  in 
suggesting  possible  means  for  their  control.  Doubtless  at  the  present 
time  most  of  the  major  sources  of  error  are  recognized.  But  the  true 
extent  of  the  errors  and  the  actual  practical  value  of  the  methods  of 
counteracting  them  can  be  determined  only  by  numerous  investiga- 
tions of  experimental  methods  under  different  conditions. 

The  present  paper  is  concerned  with  experimental  error  and  field 
plot  technic  in  preliminary  variety  and  strain  tests  with  the  small 
grains.  The  same  type  of  test  is  extensively  used  in  small  grain  im- 
provement, not  only  in  the  preliminary  testing  of  varieties,  but  also 
in  the  comparison  of  strains  and  selections.  Although  the  small  plot 
test  is  particularly  subject  to  errors  of  certain  sorts,  it  has  a decided 
advantage  over  tests  in  larger  plots  in  the  possibility  of  extensive 
replication,  which  is  probably  the  greatest  single  factor  in  the  reduc- 
tion of  experimental  error.  It  should  be  possible,  consequently,  to 
obtain  extremely  accurate  results  in  small  plot  tests  without  the  use 
of  large  experimental  areas,  when  the  errors  peculiar  to  the  small 
plot  are  understood  and  controlled. 


*Also  submitted  as  a thesis  in  partial  fulfilment  of  the  requirements  for  the  degree  of 
Doctor  of  Philosophy. 


(5) 


6 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


THE  PROBLEM. 

At  present  the  type  of  plot  most  commonly  used  for  the  pre- 
liminary testing  of  small  grain  varieties  and  strains  is  probably  the 
“rod-row.”  The  methods  of  conducting  rod-row  tests  described  by 
Love  and  Craig8  may  be  considered  typical.  The  varieties  or  strains 
are  sown  by  hand  in  rows  one  foot  apart,  usually  opened  and  covered 
with  a wheel  hoe  or  similar  implement.  The  seed  for  each  row  is 
weighed  out  in  a quantity  equivalent  to  ordinary  rates  of  seeding  in 
field  practice.  In  harvesting,  six  inches  or  a foot  at  the  end  of  the 
row  is  discarded,  to  prevent  increase  in  yield  by  reason  of  the  more 
favorable  space  conditions  at  the  ends  of  the  rows.  The  list  of  va- 
rieties is  repeated  in  several  series,  and  the  results  averaged  to  reduce 
the  error  from  plot  variability.  A check  variety  is  grown  in  every 
tenth  row  to  indicate  the  variability  of  the  field. 

The  use  of  rod-row  tests  involves  several  errors,  derived  principally 
from  the  modified  conditions  under  which  the  plants  are  grown.  The 
object  of  the  test  is  to  discover  the  relative  value  of  the  strains  under 
field  conditions,  and  therefore  any  modification  of  field  conditions 
which  may  favor  some  sorts  more  than  others  introduces  error.  The 
wide  spacing  between  rows,  with  consequently  heavier  seeding  in  the 
row  for  any  given  rate  of  planting;  the  hand  seeding  and  covering,  re- 
sulting usually  in  slightly  ridged  rather  than  slightly  furrowed  rows; 
and  the  growing  of  different  varieties  in  single  rows,  in  competition 
with  other  varieties  rather  than  with  their  own  kind,  are  examples  of 
typical  conditions  which  may  be  expected  to  favor  some  varieties  more 
than  others.  Consequently  the  best  varieties  in  the  rod-row  test  are 
not  necessarily  the  best  varieties  under  field  culture,  even  when  soil  and 
seasonal  variability  are  reduced  to  the  minimum  by  replication  of  plots 
and  repetition  of  the  test  through  a series  of  seasons. 

Such  sources  of  error  as  those  mentioned  do  not  necessarily  affect 
the  variability  of  the  yields  of  replicate  plots,  as  Kiesselbach5  has 
pointed  out,  and  are  therefore  more  likely  to  escape  notice.  They  are 
systematic  errors  affecting  the  yields  of  replicate  plots  similarly. 
Marked  superiority  of  Turkey  wheat  over  Fulcaster  in  a variety  test 
in  Kansas  does  not  indicate  the  superiority  of  Turkey  over  Fulcaster 
in  Illinois,  no  matter  how  low  plot  variability  in  the  variety  test  may 
be,  because  the  growing  conditions  in  Illinois  are  different  from  the 
growing  conditions  in  Kansas.  Similarly  the  superiority  of  Turkey 
wheat  over  Fulcaster  in  a rod-row  test  may  not  mean  its  superiority 
under  field  conditions  in  the  same  locality,  because  here  again  growing 


Experiments  in  Field  Plot  Technic 


7 


conditions  are  different.  The  error  in  applying  the  results,  though  of 
course  much  less  in  degree,  is  similar  in  kind.  And,  since  the  rod-row 
test  has  no  purpose  but  to  indicate  the  relative  value  of  the  strains 
tested,  for  field  conditions,  any  pronounced  tendency  to  favor  some 
varieties  at  the  expense  of  others  is  fatal  to  its  object. 

Ordinarily,  however,  the  rod-row  test  is  only  the  first  stage  in 
variety  testing,  and  final  recommendations  are  based  upon  results  of 
tests  under  conditions  which  approach  those  of  field  culture  more 
closely.  When  the  elimination  of  varieties  in  the  rod-row  tests  is 
not  extremely  strict  a considerable  latitude  may  be  allowed,  and  under 
these  conditions  the  rod-row  test  has  served  a valuable  purpose.  It 
is  of  course  desirable  nevertheless  to  reduce  these  errors  to  the  greatest 
possible  extent. 

Probably  the  most  important  of  the  errors  mentioned  is  that  arising 
from  the  competition  between  different  varieties,  in  the  single-row  test. 
Obviously  a variety  grown  in  a single  row  between  two  different  va- 
rieties may  yield  considerably  more  or  less  than  the  same  variety 
grown  between  two  rows  of  its  own  kind.  Various  expedients  for  re- 
ducing varietal  competition  have  been  suggested.  Sometimes  the  order 
of  varieties  is  changed  in  each  series  to  bring  together  different  va- 
rieties and  thus  tend  to  equalize  the  effects  of  competition ; sometimes 
an  attempt  is  made  to  grow  the  varieties  in  such  order  as  to  bring 
together  those  of  similar  habit,  and  thus  to  reduce  the  effects  of 
competition.  Probably  the  most  effective  method  is  to  grow  border 
rows  which  may  be  discarded,  and  some  investigators  therefore  use 
three-row  or  five-row  blocks,  in  which  the  outer  row  on  each  side  is 
discarded. 

The  principal  objection  to  the  use  of  border  rows  in  the  increased 
area  required  to  test  the  same  number  of  strains,  and  the  large  pro- 
portion of  the  crop  which  is  not  harvested  for  yield.  This  is  par- 
ticularly true  when  3-row  blocks  are  used,  since  in  this  case  two- 
thirds  of  the  field  is  used  for  border  protection.  The  border  rows  may 
be  used  for  seed,  but  two-thirds  of  the  field  is  of  course  much  more 
than  is  required  ordinarily  for  this  purpose.  When  5-row  blocks  are 
used  the  proportion  of  the  crop  harvested  for  yield  is  increased  from 
one-third  to  three-fifths,  though  it  is  an  increase  in  size  of  plot,  with 
some  decrease  in  replication,  so  that  there  may  be  no  gain  in  accuracy. 
There  is  a possibility  that  the  effect  of  competition  on  the  yield  of  5-row 
blocks  may  be  slight  enough  to  permit  the  harvesting  of  all  five  rows 
for  yield,  particularly  if  the  varieties  may  be  effectively  arranged  for 
the  reduction  of  competition.  At  any  rate,  in  such  plots  the  error 
from  competition  may  be  expected  to  be  much  less  than  that  in  single- 


8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


row  plots,  since  only  two  of  the  five  rows  are  subject  to  competition 
with  a different  variety,  and  each  of  these  is  subject  to  such  compe- 
tition on  only  one  instead  of  on  both  sides. 

Another  phase  of  the  question  which  should  not  be  overlooked 
is  the  effect  of  adding  border  rows  on  the  error  from  soil  variability. 
If,  for  example,  each  rod-row  is  to  be  protected  from  competition  by 
two  border  rows,  the  test  will  require  three  times  as  large  a field  as 
the  same  test  without  the  border  rows.  This  can  hardly  fail  to  in- 
crease materially  the  variability  of  the  yields  of  replicate  plots,  to  an 
extent  which  will  vary  with  the  uniformity  of  the  field  concerned. 
The  use  of  border  rows  may  thus  necessitate  the  use  of  an  even  greater 
number  of  replications  for  the  same  degree  of  accuracy,  as  far  as 
plot  variability  is  concerned.  It  is  possible  that  3-row  plots  (whether 
or  not  provided  with  border  rows)  may  require  less  replications 
for  a given  degree  of  accuracy  than  single-row  plots,  on  account  of 
their  larger  size.  It  is  possible  also  that  5-row  plots,  because  of 
their  size,  may  have  an  advantage  over  3-row  plots  in  reducing  va- 
riability, great  enough  to  justify  in  practice  harvesting  all  five  rows 
for  yield,  rather  than  harvesting  the  interior  three  rows  and  discarding 
the  border  rows. 

The  importance  of  any  practice  that  will  reduce  the  variability  of 
the  replicate  plots  is  thus  increased  when  border  rows  are  introduced. 
A familiar  method  for  this  purpose  is  the  adjustment  of  yields  by 
means  of  distributed  check  plots.  In  following  this  method  the  yields 
of  check  plots  are  considered  measures  of  the  productivity  of  the 
soil,  which  is  usually  assumed  to  vary  uniformly  between  them.  The 
yields  of  the  experimental  plots  are  adjusted  on  the  basis  of  uniform 
productivity  of  the  field  as  a whole.  Of  late  this  method  has  rather 
lost  favor  among  agronomists.  In  some  cases  the  adjustment  actually 
increases  rather  than  decreases  the  variability  of  the  replicate  ex- 
perimental plots.  Check  plots  have  not  been  used  extensively  in  ad- 
justing yields  in  rod-row  tests,  principally  because  of  the  great  in- 
crease in  computation  necessary  in  adjusting  the  yields  of  such  a large 
number  of  plots. 


Experiments  in  Field  Plot  Technic 


9 


PLAN  AND  METHOD  OF  INVESTIGATION 

The  experiments  here  reported  were  designed  to  obtain  informa- 
tion on  several  factors  affecting  the  accuracy  of  preliminary  variety 
and  strain  tests,  with  a view  to  devising,  if  possible,  an  improved 
technic  for  this  important  phase  of  crop  improvement  work.  The 
data  obtained  bear  directly  on  the  following  points : 

1.  The  extent  of  error  from  varietal  competition  in  bor- 
der rows,  and  the  relation  of  such  competition  to  the  charac- 
teristics of  the  varieties, 

2.  The  relative  variability  of  plots  of  1,  3,  and  5 rows,  and 
the  number  of  replications  necessary  for  a given  degree  of 
precision  with  plots  of  the  three  sizes,  and 

3.  The  effect  on  variability  of  adjusting  yields  by  means 
of  check  plots. 

Terminology. — In  this  report  the  term  plot  will  be  used  to  des- 
ignate an  area  on  which  a single  variety  or  strain  is  grown,  in  com- 
parison with  other  varieties  or  strains,  in  other  plots.  The  plot  may 
consist  of  one  or  more  rows.  A plot  of  more  than  one  row  may  also  be 
referred  to  as  a block.  The  single  outside  rows  of  the  block  are  the 
border  rows.  A single-row  plot  protected  from  competition  by  border 
rows,  which  are  to  be  discarded,  will  be  spoken  of  as  a protected 
single-row  plot.  A protected  single-row  plot  is  therefore  a 3-row  plot 
with  border  rows  discarded,  and  a protected  3-row  plot  is  a 5-row  plot 
with  border  rows  discarded.  The  phrase  “3-row  plots  replicated  five 
times”  will  be  used  to  refer  to  3-row  plots  in  five  systematically  dis- 
tributed locations,  not  in  six.  The  area  on  which  a complete  variety 
or  strain  test  is  conducted  is  spoken  of  as  an  experiment  field,  or  simply 
a field.  A group  of  plots  including  one  plot  of  each  variety  or  strain 
tested  is  a series.  When  four  replications  are  used  there  are  four 
series  of  plots.  The  group  of  contiguous  plots  from  one  side  of  the 
field  to  the  other  constitutes  a range.  The  ranges  are  separated  by 
alleys. 

Thus  the  field  shown  in  figure  1 consists  of  sixteen  ranges,  each 
range  including  twenty-nine  5-row  (or  protected  3-row)  plots.  Ninety- 
six  varieties  were  tested  on  this  field,  each  replicated  four  times. 
Ranges  I to  IV,  inclusive,  make  up  the  first  series,  V to  VIII  the  sec- 
ond, IX  to  XII  the  third,  and  XIII  to  XVI  the  fourth.  Each  of  the 
four  strips  running  lengthwise  of  the  field  and  separated  by  the  check 
plots  may  also  be  considered  a series. 

All  yields  are  expressed  in  bushels  per  acre  by  weight,  computed 
on  the  basis  of  60  pounds  per  bushel  for  wheat,  48  pounds  for  barley, 


10 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


B 

CK 

I 

17 

33 

43 

65 

81 

CK 

5 

2/ 

37 

53 

69 

65 

CK 

9 

25 

41 

57 

73 

89 

CK 

ft 

29 

45 

61 

77 

93 

CK 

B 

CK 

2 

18 

34 

50 

66 

62 

CK 

6 

22 

38 

54 

70 

86 

CK 

10 

26 

42 

58 

74 

90 

CK 

14 

30 

46 

62 

7 8 

94 

CK 

8 

CK 

3 

IS 

35 

5/ 

67 

83 

CK 

7 

23 

3 9 

55 

7/ 

87 

CK 

II 

27 

43 

59 

75 

91 

CK 

15 

31 

47 

63 

79 

95 

CK 

B 

CK 

4 

20 

36 

52 

68 

64 

CK 

8 

24 

40 

56 

72 

63 

CK 

12 

28 

44 

60 

76 

92 

CK 

16 

32 

48 

64 

80 

96 

CK 

B 

CK 

5 

2/ 

37 

53 

69 

85 

CK 

9 

25 

41 

57 

73 

83 

CK 

13 

29 

45 

61 

77 

93 

CK 

17 

33 

49 

65 

8/ 

CK 

B 

CK 

6 

22 

33 

54 

70 

86 

CK 

10 

26 

42 

58 

74 

90 

CK 

IK 

30 

46 

62 

78 

94 

CK 

2 

18 

34 

50 

66 

82 

CK 

B 

CK 

7 

23 

39 

55 

71 

87 

CK 

// 

27 

43 

59 

75 

91 

CK 

15 

31 

47 

63 

79 

95 

CK 

3 

19 

35 

51 

67 

83 

CK 

B 

CK 

<9 

24 

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56 

72 

88 

CK 

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28 

44 

60 

76 

92 

CK 

J6 

32 

4 8 

64 

80 

96 

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57 

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89 

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13 

29 

45 

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77 

93 

CK 

I 

17 

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49 

65 

8! 

CK 

5 

21 

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53 

69 

85 

CK 

B 

CK 

10 

26 

42 

58 

74 

SO 

CK 

/4 

30 

4 6 

62 

78 

94 

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2 

18 

34 

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66 

82 

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6 

22 

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86 

CK 

B 

CK 

// 

21 

43 

59 

75 

SI 

CK 

15 

31 

47 

63 

79 

95 

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3 

13 

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83 

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7 

23 

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55 

71 

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CK 

B 

CK 

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28 

44 

60 

76 

92 

CK 

16 

32 

46 

64 

80 

96 

CK 

4 

20 

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68 

84 

CK 

6 

24 

40 

56 

72 

86 

CK 

B 

CK 

ft 

29 

45 

61 

77 

93 

CK 

/ 

17 

33 

45 

65 

81 

CK 

5 

21 

37 

53 

69 

85 

CK 

9 

25 

41 

57 

73 

89 

CK 

B 

CK 

H 

30 

46 

62 

78 

94 

CK 

2 

18 

34 

50 

66 

82 

CK 

6 

22 

38 

54 

70 

86 

CK 

10 

26 

42 

58 

74 

90 

CK 

B 

CK 

15 

31 

47 

63 

73 

95 

CK 

3 

19 

35 

5/ 

67 

83 

CK 

7 

23 

39 

55 

7/ 

87 

CK 

II 

27 

43 

59 

75 

9/ 

CK 

B 

CK 

16 

32 

43 

64 

00 

96 

CK 

4 

20 

36 

52 

66 

m 

CK 

8 

24 

40 

56 

72 

88 

CK 

12 

28 

44 

60 

76 

92 

CK 

Figure  1.— Planting  Plan  of  Wheat  Variety  Tests  1920  and  1921. 
Legend:  B,  border.  CK,  check.  Numbers  1-96,  planting  numbers  of  varieties 

tested  as  given  in  Tables  4 and  5. 


Experiments  in  Field  Plot  Technic 


11 


and  32  pounds  for  oats.  The  measures  of  variability  used  are  the 
average  deviation,  the  standard  deviation,  and  the  probable  error. 
These  were  computed  according  to  the  following  formulae: 


n 


E=±.6745  cr . 


in  which  A.D.  = average  deviation,  o-=standard  deviation,  E = prob- 
able error  (of  a single  determination),  d = the  deviation  of  a single 
variate  from  the  mean,  and  n = the  number  of  variates.  The  correla- 
tion coefficient  r was  determined  by  the  formula 


/S(dxdy) 


and  the  probable  error  of  the  correlation  coefficient  Er  by  the  formula 


Er 


.6745  (1  — r2) 

V" 


The  tests  reported  are  of  two  kinds,  variety  tests  and  strain  tests. 
The  variety  tests  were  comparisons  of  commercial  varieties,  most 
of  which  were  taxonomically  distinct.  A number  of  pure  line  selec- 
tions were  included  in  the  wheat  variety  tests.  The  strain  tests  were 
comparisons  of  a considerable  number  of  commercial  lots  of  the  same 
variety  obtained  from  different  sources.  These  strains,  so-called  for 
convenience,  are  not,  except  in  a very  few  cases,  pure  lines.  Some  of 
them  are  possibly  identical,  and  all  the  strains  of  any  one  variety  are 
of  course  very  similar,  since  they  are  taxonomically  the  same. 

Procedure. — In  the  seasons  of  1919,  1920,  1921,  tests  of  va- 
rieties and  strains  of  oats,  barley,  and  wheat  were  conducted  in  blocks 
consisting  of  five  rows  ten  inches  apart  and  usually  18  feet  long. 
From  24  to  96  varieties  were  included  in  each  test,  and  from  three  to 
six  (usually  four)  replications  were  used.  The  planting  order  in  each 
case  was  designed  on  a plan  similar  to  that  illustrated  in  figure  1. 
It  will  be  noted  that  the  check  plots  were  in  continuous  strips,  that 
each  variety  was  represented  in  each  quarter  of  the  field,  whether 
divided  from  east  to  west  or  from  north  to  south,  and  that  in  all 
four  series  each  variety  occupied  the  same  position  with  relation  to 
the  check  plots,  and  had  the  same  varieties  adjoining  it  on  either  side. 


12 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


The  rows  in  some  cases  ran  east  and  west,  and  in  some  cases  north 
and  south. 

All  these  plots  were  seeded  with  a 5-row  nursery  drill,  built  from 
plans  furnished  by  Professor  T.  A.  Kiesselbach  of  the  Nebraska  Sta- 
tion. This  is  a hoe  drill  designed  for  rapid  and  thorough  cleaning 
between  plots.  Photographs  of  it  have  been  published  in  reports  of 
earlier  work  on  field  plot  technic  at  the  Nebraska  Station  (Mont- 
gomery14 page  57,  and  Kiesselbach  5 page  16).  Its  use  resulted  in  uni- 
form seeding  and  covering  and  accurate  spacing  between  rows,  with  a 
close  approach  to  ordinary  field  conditions  in  the  state  in  which  the 
field  was  left  after  seeding.  Each  field  was  seeded  in  a single  day. 

All  plots  were  harvested  by  hand  with  sickles,  a foot  at  each 
end  of  each  row  discarded,  and  the  remainder  (usually  16  feet)  tied 
in  a bundle  and  hung  in  a ventilated  shed  to  dry.  In  1919  and  1920 
each  row  was  bundled  and  threshed  separately;  in  1921  the  border 
rows  of  each  5-row  block  were  bundled  separately  and  the  three  in- 
terior rows  bundled  together.  Yields  were  determined  by  weighing 
in  grams  at  the  time  of  threshing.  All  final  yields  were  converted 
to  bushels  per  acre  and  are  so  expressed. 

Work  of  1919. — In  1919  tests  were  conducted  with  barley  and 
oats.  Thirty  varieties  of  barley  were  grown,  each  in  3 replicate  plots. 
The  test  comprised  three  ranges  of  185  rows  each,  including  21  check 
plots,  or  one  in  every  sixth  plot.  The  barley  was  drilled  at  the  rate 
of  eight  pecks  per  acre,  on  March  21,  in  rows  running  north  and  south. 
The  rows  were  14  feet  long  and  10  inches  apart.  They  were  cut  to  12 
feet  in  harvesting.  The  planting  plan  is  shown  in  figure  2.  Conditions 


CK 

/ 

2 

3 

9 

5 

cn 

6 

7 

8 

9 

10 

OH 

II 

12 

13 

19 

15 

a 

16 

17 

18 

19 

20 

CK 

21 

22 

23 

29 

25 

CK 

26 

27 

28 

29 

30 

CK 

B 

CK 

2/ 

22 

23 

2i 

25 

cn 

26 

21 

28 

29 

30 

CH 

1 

2 

3 

9 

5 

cn 

6 

7 

8 

9 

10 

CH 

11 

12 

13 

19 

15 

CH 

16 

n 

!8 

19 

20 

CK 

B 

CK 

// 

12 

/3 

M 

15 

CK 

16 

17 

18 

!9 

20 

CK 

2/ 

22 

23 

29 

25 

CK 

26 

27 

28 

29 

30 

GK 

/ 

2 

3 

9 

5 

CH 

6 

7 

8 

9 

to 

CH 

B 

Figure  2. — Planting  Plan  of  Barley  Variety  Test  1919.  Legend:  B, 

border.  CK,  check.  Numbers  1-30,  planting  numbers  of  varieties  tested,  as 
given  in  Table  1. 

were  fairly  favorable,  and  the  yields  of  the  adapted  varieties  were 
slightly  higher  than  the  average  obtained  under  the  conditions  at  Co- 
lumbia. Two  varieties,  Italian  and  Australian  White,  gave  extremely 
low  yields  and  were  excluded.  Another,  Sandrel,  was  represented 
only  in  two  series,  and  was  also  excluded.  The  yields  of  the  remain- 


Experiments  in  Field  Plot  Technic 


13 


in g 27  varieties  are  shown  in  Table  1.  The  planting  numbers  given  in 
this  table  correspond  to  those  shown  in  the  diagram  of  the  field  (figure 
2.) 


Table  1. — Yields  oe  Barley  Varieties. 
In  Bushels  per  Acre.  1919. 


Planting 

number 

Variety 

Average  Yield 
3 interior  rows  5 rows 

1 

Hanna  906 

12.55 

12.57 

2 

Steigum  907 

19.90 

19.65 

3 

Luth  908 

23.65 

23.40 

4 

Eagle  913 

20.40 

20.13 

5 

Italian  914* 

6.70 

6.57 

6 

Servian  915 

19.85 

19.86 

7 

Odessa  916 

13.75 

13.41 

8 

Lion  923 

21.75 

22.14 

9 

Australian  White  925* 

1.45 

1.74 

10 

Horn  926 

21.25 

21.54 

11 

Odessa  927 

20.80 

19.53 

12 

Summit  929 

23.05 

24.03 

13 

Mariout  932 

18.75 

18.15 

14 

Odessa  934 

10.30 

9.84 

15 

Peruvian  935 

22.25 

20.55 

16 

Trebi  936 

30.90 

30.96 

17 

Sandrel  937* 

35.90 

33.48 

18 

Oderbrucker  940 

23.35 

23.79 

19 

Frankish  953 

22.50 

22.05 

20 

Manchuria  956 

30.80 

30.03 

21 

Oderbrucker  957 

29.45 

29.52 

22 

Manchuria  x Champion  of  Vermont  959 

18.30 

17.49 

23 

Luth  972 

25.05 

26.28 

24 

Red  River  973 

27.25 

28.14 

25 

Featherston  1118 

28.25 

27.00 

26 

Featherston  1119 

25.80 

25.83 

27 

Featherston  1120 

34.35 

35.49 

28 

Hanna  x Champion  of  Vermont  1121 

13.75 

13.92 

29 

Manchuria  1125 

20.35 

20.94 

30 

Malting  1129 

17.25 

16.44 

Mean 

22.06 

21.95 

Forty  varieties  of  oats  were  compared  in  1919,  but  only  24  of 
these  could  be  replicated  4 times  and  the  remaining  16  were  duplicated. 
The  planting  plan  was  therefore  arranged  as  for  32  varieties,  and 
these  16  varieties  grown  in  two  plots  each  in  place  of  eight  varieties 


?Hi  and  .A1u.stralia?  White  925  were  omitted  from  all  computations  because  of 
their  extremely  low  yields,  and  Sandrel  937  because  omitted  in  the  third  series. 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


CK 

/ 

2 

3 

4 

5 

6 

7 

3 

CK 

9 

10 

n 

12 

13 

14 

15 

16 

CK 

17 

16 

19 

20 

2/ 

22 

23 

24 

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25 

26 

27 

26 

89 

30 

31 

32 

CH 

B 

CK 

33 

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33 

36 

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39 

40 

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n 

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CK 

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CK 

17 

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28 

29 

30 

31 

32 

CK 

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CK 

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CK 

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17 

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20 

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33 

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35 

36 

37 

36 

39 

40 

CK 

/ 

2 

3 

4 

5 

6 

7 

6 

CK 

B 

CK 

5 

E 

E 

D 

n 

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L 

T 

/ 

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L 

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C 

8 

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I 

0 

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B 

CK 

/ 

2 

3 

4 

5 

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6 

7 

8 

9 

10 

CK 

II 

12 

13 

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15 

CK 

/ 

2 

3 

4 

5 

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6 

7 

8 

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70 

CK 

II 

18 

13 

14 

15 

CK 

B 

CK 

// 

12 

13 

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15 

CK 

I 

2 

3 

4 

5 

CK 

6 

7 

6 

9 

10 

CK 

II 

12 

13 

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15 

CK 

/ 

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3 

4 

5 

CK 

6 

X 

X 

9 

10 

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CK 

6 

7 

8 

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12 

13 

14 

15 

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/ 

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4 

5 

CK 

6 

7 

8 

9 

10 

CK 

// 

12 

13 

14 

15 

CK 

X 

Z 

X 

X 

X 

CK 

B 

Figure  3. — Planting  Plan  oe  Oats  Variety  and  Strain  Tests  1919. 
Legend:  B,  border.  CK,  check.  Numbers  1-40  in  first  four  ranges,  planting 

numbers  of  oats  varieties,  as  given  in  Table  2.  Numbers  1-15  in  last  three 
ranges,  planting  numbers  of  oats  strains,  as  given  in  Table  3.  X,  test  plots 
planted  to  check  variety  because  of  insufficient  supply  of  seed. 


Table  2. — Yields  of  Oats  Varieties. 
In  Bushels  per  Acre.  1919. 


Planting 

number 

Variety 

Average  yield 
Four  series 

in  interior  rows 
Three  series 

1 

A.  Sterilis  nigra 

30.0 

31.7 

2 

Black  Mesdag 

44.2 

44.7 

3 

C.  I.  602 

35.4 

38.1 

4 

C.  I.  603 

53.9 

55.1 

5 

C.  I.  620 

13.1 

14.1 

6 

Early  Champion 

55.5 

53.9 

7 

Early  Gothland 

54.1 

52.8 

8 

Garton  473 

30.6 

31.7 

9 

Garton  585 

21.7 

23.0 

10 

Golden  Giant 

42.0 

44.9 

11 

Irish  Victor 

69.6 

70.2 

12 

Japan  Selection 

47.9 

50.9 

13 

June 

43.1 

44.5 

14 

Kherson  Selection 

67.2 

63.1 

15 

Fulghum  042 

60.9 

57.1 

16 

Lincoln 

51.5 

50.3 

17 

Monarch 

56.0 

53.4 

18 

North  Finnish 

51.0 

49.5 

19 

Scottish  Chief 

59.3 

60.1 

20 

Sparrow  bill  (Missouri) 

39.8 

41.3 

21 

Sparrow  bill  (Cornell) 

42.3 

45.7 

22 

Tobolsk  1 

52.6 

57.3 

23 

Tobolsk  2 

46.1 

51.9 

24 

White  Tartar 

49.7 

50.3 

Mean 

46.6 

47.3 

Experiments  in  Fieed  Plot  Technic 


15 


in  four  plots  each,  as  shown  in  figure  3.  The  rows  were  14  feet  long 
and  were  cut  to  12  feet  in  harvesting.  This  is  a convenient  size  of 
plot  for  oats  tests  with  10  inches  distance  between  rows,  when  the 
border  rows  are  discarded,  since  the  total  yield  of  three  rows  in 
grams,  divided  by  10,  gives  the  yield  in  bushels  per  acre.  The  oats 
were  planted  at  the  rate  of  10  pecks  per  acre,  on  March  18,  in  rows 
running  north  and  south.  The  season  was  favorable  and  a good  yield 
of  the  better  varieties  was  obtained.  The  yields  of  the  24  varieties 
replicated  four  times  are  shown  in  Table  2. 

The  oats  strain  test  was  conducted  on  the  same  field,  as  shown  in 
figure  3,  directly  south  of  the  oats  variety  test.  In  planting,  these  two 
tests  were  handled  as  one ; and  the  rate,  date,  and  method  of  planting 
were  the  same.  The  strains  tested  were  15  strains  of  oats  obtained 
under  the  name  Red  Rustproof  from  various  experiment  stations  and 
seedsmen.  Three  of  these  strains,  0121,  0124,  and  0127,  were  not  true 
to  name,  but  the  remainder  were  taxonomically  Red  Rustproof  oats, 
as  described  by  Etheridge2.  The  oats  strains  were  tested  in  six  series, 
with  check  plots  in  every  sixth  plot.  The  line  of  check  plots  on  the  west, 
however,  gave  abnormally  low  yields,  probably  because  they  were 
located  partly  on  a dead  furrow  at  the  edge  of  the  experiment  field. 
On  account  of  shortage  of  seed  some  of  the  varieties  could  not  be 
planted  in  the  last  series.  The  first  and  last  series  were  therefore  dis- 


Table  3. — Yields  of  Oats  Strains  (Red  Rustproof). 
In  Bushels  per  Acre.  1919. 


Planting 

number 

Accession 

number 

Average  yield 

3 interior  Rows  5 Rows 

1 

0119 

49.58 

49.41 

2 

0120 

45.83 

44.51 

3 

0121* 

49.43 

53.01 

4 

0122 

47.85 

49.59 

5 

0123 

53.55 

53.47 

6 

0125 

50.18 

49.19 

7 

0126 

44.85 

45.81 

8 

0127* 

38.55 

36.67 

9 

0124* 

63.90 

67.46 

10 

0133 

48.00 

46.49 

11 

0128 

53.55- 

53.15 

12 

0129 

49.35 

49.01 

13 

0130 

52.73 

51.89 

14 

0131 

48.60 

47.84 

15 

0132 

55.13 

55.44 

Mean 

50.07 

50.20 

#Not  taxonomically  Red  Rustproof. 


16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


carded.  The  average  yields  of  the  15  strains  in  the  four  remaining 
series  are  shown  in  Table  3. 

Work  of  1920. — Wheat  varieties  were  grown  in  5-row  blocks  in 
1919-20.  Ninety-six  varieties  were  included  in  the  test,  four  replica- 
tions being  used.  Fultz  wheat  was  grown  as  a check  in  every  sixth 
plot.  The  rows  were  18  feet  long  and  were  cut  to  16  feet  in  harvest- 
ing. The  direction  of  the  rows  was  east  and  west.  The  planting  plan 
is  shown  in  figure  1.  The  wheat  was  sown  October  15,  at  the  rate  of 
6 pecks  per  acre.  There  was  considerable  winter  injury  in  the  plots 
and  the  condition  of  the  wheat  in  early  spring  was  rather  poor.  The 
yields  obtained  are  shown  in  Table  4. 


Table  4. — Yields  oe  Wheat  Varieties. 
In  Bushels  per  Acre  1980. 


Average  yield  - Average  yield 

Planting  3 Interior  5 Planting  3 Interior  5 


number  Variety 

Rows 

Rows 

number  Variety 

Rows 

Rows 

1 Beechwood  Hybrid  No.  12.. 

10.8 

11.1 

50 

Michigan  Wonder 

No. 

141  . 

. 10.7 

10.3 

2 Beechwood  Hybrid  No.  81.. 

12.8 

14.1 

51 

Michigan  Wonder 

No. 

155 

. 10.1 

9.8 

3 Beechwood  Hybrid  No.  85.. 

12.5 

12.7 

52 

Michigan  Wonder 

No. 

209 

. 12.1 

12.5 

4 Beechwood  Hybrid  No.  87.. 

14.2 

13.7 

53 

Michigan  Wonder 

No. 

211 

. 9.9 

9.9 

5 Beechwood  Hybrid  No.  202. 

11.9 

12.5 

54 

Michigan  Wonder 

No. 

221 

. 11.0 

11.1 

6 Beechwood  Hybrid  No.  207. 

13.2 

13.6 

55 

New  York  123-32  . . 

. 17.2 

17.5 

7 C.  I.  3808  

16.2 

16.6 

56 

N iagara  

. 13.8 

13.5 

8 C.  I.  3846  

14.2 

15.6 

57 

Nigger  

. 11.8 

11.8 

9 C.  I.  3972  

14.7 

15.6 

58 

Old  Ironclad  . . . 

. 12.5 

13.2 

10  C.  I.  3980  

16.4 

17.5 

59 

Poole  

. 10.5 

10.7 

11  C.  I.  3988  

16.7 

17.0 

60 

Poole  No.  3 . . . . 

. 11.7 

11.0 

12  C.  I.  4004  

14.3 

14.0 

61 

Poole  B-3  

. 12.5 

13.3 

13  Common  Rye  

17.3 

18.5 

62 

Portage  

. 15.9 

17.3 

14  Dawson’s  Golden  Chaff  .... 

13.0 

12.3 

63 

Pride  of  Indiana 

. 14.2 

14.4 

15  Deitz  

15.1 

14.6 

64 

Pride  of  Genessee  . . 

. 15.7 

18.1 

16  Early  Ripe  

12.2 

12.6 

65 

Reliable  

. 12.6 

12.9 

17  Early  Ripe  No.  26  

13.2 

14.0 

66 

Red  Cross  

. 13.1 

13.1 

18  Early  Red  Clawson  

9.9 

9.5 

67 

Red  May  

. 14.8 

14.8 

19  Farmer’s  Friend  

18.8 

19.9 

68 

Red  Rock  (Indiana) 

. 18.7 

19.7 

20  Fulcaster  

14.4 

15.3 

69 

Red  Rock  (Michigan) 

. 7.5 

6.8 

21  Fultz  GArchias)  

12.2 

13.0 

70 

Red  Wave  

. 12.9 

12.7 

22  Gold  Coin  

11.7 

12.2 

71 

Rochester  Red  . . 

. 12.7 

12.9 

23  Greene  County  

15.6 

15.1 

72 

Rosen  Rye  

. 20.7 

24.0 

24  Harvest  King  No.  7 

13.4 

14.2 

73 

S.  P.  I.  11616  . 

. 10.3 

10.9 

25  Harvest  Queen  

9.6 

9.8 

74 

S.  P.  I.  26012  . 

. 13.4 

12.9 

26  Hickman  

T.  8 

8(2 

75 

S.  P.  I.  26013  . 

15.2 

15.5 

27  Illini  Chief  

17.5 

18.7 

76 

S.  P.  I.  26014  . 

. 17.5 

18.8 

28  Jones  Climax  

19.1 

20.7 

77 

S.  P.  I,  26015  . 

. 13.2 

13.4 

29  Kanred  

21.0 

22.7 

78 

S.  P.  I.  26017  . 

. 13.4 

13.5 

30  Kessinger  

18.0 

19.3 

79 

S.  P.  I.  26018  . 

. 13.6 

13.6 

31  Kharkov  

18.9 

20.1 

80 

S.  P.  I.  26019  . 

. 11.6 

11.6 

32  Leap’s  Prolific  

14.2 

14.8 

81 

S.  P.  I.  26022  . 

10.6 

10.1 

33  Mediterranean  No.  8 

9.1 

9.5 

82 

S.  P.  I.  26023  . 

. 9.1 

8.5 

34  Michigan  Amber  

10.7 

11.3 

83 

S.  P.  I.  26025  . 

. 12.3 

13.1 

35  Michigan  Amber  (Indiana) 

17.0 

17.9 

84 

S.  P.  I.  26029  . 

. 15.4 

15.6 

36  Michigan  Amber  No.  7 ... 

10.5 

10.8 

85 

S.  P.  I.  26085  . 

. 13.2 

13.3 

37  Michigan  Amber  No.  12  ... 

9.3 

9.3 

86 

Treadwell  

. 12.7 

12.8 

38  Michigan  Wonder  

10.9 

11.1 

87 

Valley  

. 12.4 

12.1 

39  Michigan  Wonder  No.  4 ... 

12.4 

12.7 

88 

Velvet  Chaff  No. 

2 . 

. 14.1 

12.8 

40  Michigan  Wonder  No.  8 ... 

10.8 

11.0 

89 

Velvet  Chaff  No. 

8 .. 

. 9.0 

9.3 

41  Michigan  Wonder  No.  21  . . 

8.2 

8.7 

90 

Ziegler’s  Fly  Proof  . 

. 10.9 

11.7 

42  Michigan  Wonder  No.  53  . . 

8.5 

9.6 

91 

13D-4a  

. 14.1 

13.9 

43  Michigan  Wonder  No.  54  . . 

11.3 

10.7 

92 

37a-4  

. 14.6 

14.7 

44  Michigan  Wonder  No.  83  . . 

13.6 

13.7 

93 

Fulcaster  (Co-op) 

. 17.4 

18.5 

45  Michigan  Wonder  No.  96  . . 

9.7 

9.7 

94 

Fultz  (Co-op) 

. 15.2 

15.9 

46  Michigan  Wonder  No.  103  . 

9.7 

9.1 

95 

Kanred  (Co-op)  . 

. 19.1 

20.6 

47  Michigan  Wonder  No.  116  . 

16.4 

15.8 

96 

Poole  (Co-op)  . . 

. 19.4 

21.0 

48  Michigan  Wonder  No.  130  . 

14.3 

14.2 

49  Michigan  Wonder  No.  140  . 

12.3 

12.7 

Mean  

13.4 

13.8 

Experiments  in  Fieed  Plot  Technic 


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18 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


Work  of  1921. — In  1920-21  ninety-six  varieties  of  wheat 
were  again  tested  by  this  method.  Many  of  the  varieties  were  the  same 
as  those  tested  in  the  preceding  year,  about  20  varieties  being  eliminated 
and  a corresponding  number  added.  The  planting  plan  was  the  same 
as  that  of  the  preceding  season.  Poole  wheat  was  used  as  a check 
variety.  The  wheat  was  drilled  at  the  rate  of  5 pecks  per  acre,  October 
6,  in  rows  running  east  and  west.  The  season  was  favorable,  but 
yields  were  reduced  by  the  very  rapid  ripening  of  the  wheat  caused  by 
the  hot  dry  weather  in  the  second  and  third  weeks  of  June.  The  yields 
are  shown  in  Table  5. 


Table  6. — Yields  of  Wheat  Varieties  and  Mixtures 
In  Bushels  per  Acre.  1921. 


Planting 

number 

Variety 

Average  yield 
3 Interior  Rows  5 Rows 

1 

Fulcaster 

17.3 

18.6 

2 

Harvest  Queen 

14.2 

14.5 

3 

Mixture  No.  1 (1,  2,  4,  5) 

15.9 

16.2 

4 

Michigan  Wonder 

16.8 

17.8 

5 

Nigger 

10.8 

10.8 

6 

Michigan  Wonder  No.  21 

19.8 

20.8 

7 

Michigan  Wonder  No.  54 

18.9 

19.3 

8 

Mixture  No.  2 (6,  7,  9,  10) 

20.8 

21.3 

9 

Michigan  Wonder  No.  96 

18.5 

18.9 

10 

Michigan  Wonder  No.  209 

21.7 

22.6 

11 

Beechwood  Hybrid  No.  12 

17.4 

18.8 

12 

Beechwood  Hybrid  No.  85 

16.5 

17.3 

13 

Mixture  No.  3 (11,  12,  14,  15) 

17.6 

18.4 

14 

Beechwood  Hybrid  No.  87 

19.9 

19.9 

15 

Beechwood  Hybrid  No.  207 

17.4 

17.9 

16 

Michigan  Wonder  No.  221 

18.6 

20.3 

17 

Kanred 

13.6 

13.8 

18 

Mixture  No.  4 (16,  17,  19,  20) 

17.8 

18.0 

19 

New  York  123-32 

19.6 

19.7 

20 

Red  Rock 

17.6 

17.4 

21 

Red  Hussar 

16.3 

17.8 

22 

Turkey  (Kansas) 

10.8 

10.5 

23 

Mixture  No.  5 (21,  22,  24,  25) 

15.7 

15.9 

24 

Michigan  Amber 

19.2 

19.6 

25 

Nigger 

14.1 

13.4 

26 

Fulcaster  (Co-op) 

20.4 

21.2 

27 

Fulcaster  (Outl) 

20.1 

20.6 

28 

Mixture  No.  6 (26,  27,  29,  30) 

20.1 

21.0 

29 

Fulcaster  (Blazier) 

20.6 

21.5 

30 

Fulcaster  (Cowles) 

20.6 

20.6 

Mean 

17.6 

18.2 

Experiments  in  Field  Plot  Technic 


19 


On  another  field  in  1921,  a test  of  mixtures  of  varieties  and 
strains  of  wheat  in  comparison  with  their  pure  constituents  was  con- 
ducted. Each  mixture  was  made  up  of  four  varieties  or  strains,  in 
equal  quantities  of  seed  by  weight.  The  composition  of  the  mixtures 
and  the  yields  obtained  are  shown  in  Table  6.  The  planting  plan  is 
shown  in  figure  4.  This  wheat  was  drilled  at  the  rate  of  5 pecks  per 


B 

Cli 

2 

3 

4 

5 

CK 

26 

27 

26 

29 

30 

CK 

16 

n 

Id 

19 

20 

CK 

II 

12 

13 

14 

IS 

CH 

B 

B 

CH 

6 

7 

a 

9 

10 

CK 

1 

2 

3 

4- 

S 

CK 

21 

22 

23 

24 

25 

CK 

16 

17 

18 

19 

20 

CH 

B 

B 

CK 

// 

IZ 

13 

1+ 

15 

CK 

6 

r 

a 

9 

/O 

CK 

26 

27 

28 

29 

30 

CK 

2 1 

32 

23 

24 

25 

CH 

B 

B 

CK 

16 

17 

16 

19 

20 

CK 

// 

id 

13 

14 

15 

CK 

2 

3 

4 

S 

CK 

36 

37 

28 

29 

30 

CK 

B 

B 

CK 

2 1 

22 

23 

24 

25 

CK 

/6 

17 

18 

19 

20 

CK 

6 

7 

8 

9 

10 

CK 

Z 

3 

4 

S 

CK 

B 

B 

CK, 

26 

27 

26 

29 

30 

CK 

2/ 

22 

23 

24 

25 

CK 

II 

/2 

13 

14 

15 

CK 

6 

7 

8 

9 

10 

CK 

B 

Figure  4. — Planting  Plan  of  Wheat  Mixture  Test  1921.  Legend:  B, 
border.  CK,  check.  Numbers  1-30,  planting  numbers  of  varieties  and  mixtures 
tested,  as  given  in  Table  6. 

acre  in  rows  running  north  and  south,  on  October  8,  1920.  This  test 
will  be  referred  to  as  the  wheat  mixture  test. 

In  1921  tests  of  oats  varieties  and  of  oats  strains  were  also  con- 
ducted in  5-row  blocks.  Thirty-two  strains  of  Red  Rustproof,  in- 
cluding many  of  those  tested  in  1919  and  a number  of  others,  and  32 
strains  of  Kherson  oats,  obtained  in  the  same  way,  were  included  in 
the  oats  strain  test.  The  Kherson  and  Red  Rustproof  strains  were 
arranged  alternately,  and  both  Kherson  and  Red  Rustproof  checks 
were  grown,  as  shown  in  figure  5.  The  test  of  these  64  strains,  in 
four  series,  occupied  16  ranges.  The  next  eight  ranges  on  the  same 
plot  were  used  for  an  oats  variety  test  in  which  32  varieties  of  oats 
were  compared,  each  in  four  replicate  plots.  In  this  part  of  the  field 
the  Kherson  and  Red  Rustproof  check  plots  were  continued.  There 
are  thus  available  the  yields  of  120  plots  each  of  Kherson  and  Red 
Rustproof  oats,  or  five  strips  of  24  plots  of  each  arranged  in  pairs 
side  by  side.  In  both  of  these  experiments  the  rows  ran  east  and 
west,  and  were  18  feet  long,  cut  to  16  feet  in  harvesting.  The  oats 
were  drilled  on  March  12,  at  the  rate  of  10  pecks  per  acre.  The  yields 
of  oats,  particularly  of  the  later-maturing  varieties,  were  materially  re- 
duced by  the  hot  dry  weather  in  the  middle  of  June.  The  yields  of  the 
oats  varieties  are  shown  in  Table  7,  and  those  of  the  strains  in  Table  8. 


20 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


B 

74 

R 

/ 

17 

33 

49 

K 

74 

13 

29 

45 

6/ 

K 

R 

9 

25 

41 

57 

R 

R 

5 

21 

37 

53 

K 

R 

3 

B 

74 

R 

2 

18 

3.* 

50 

74 

R 

14 

30 

46 

62 

74 

R 

10 

26 

42 

S7 

74 

R 

6 

22 

38 

54 

74 

R 

B 

B 

n 

R 

3 

19 

35 

5/ 

74 

R 

15 

3/ 

47 

63 

74 

R 

// 

27 

43 

59 

R 

R 

7 

23 

39 

55 

74 

R 

B 

B 

K 

R 

f 

ZO 

36 

52 

K 

R 

76 

52 

48 

i Cb  1 

K 

R 

72 

28 

44 

60 

K 

R 

8 

24 

40 

56 

K 

R 

B 

B 

74 

R 

5 

2! 

37 

53 

74 

R 

/ 

17 

33 

49 

74 

74 

73 

29 

45 

61 

74 

R 

9 

25 

41 

57 

R 

R 

B 

B 

74 

R 

6 

2Z 

38 

54 

K 

R 

2 

te 

54 

SO 

74 

74 

J4 

30 

46 

62 

n 

R 

/ 0 

26 

42 

58 

K 

R 

3 

B 

K 

R 

7 

23 

39 

55 

K 

R 

3 

79 

35 

51 

74 

R 

15 

3/ 

47 

63 

K 

R 

// 

27 

43 

59 

K 

R 

B 

B 

74 

R 

8 

2* 

40 

56 

74 

R 

4 

20 

36 

52 

74 

R 

16 

32 

48 

64 

74 

R 

J2 

26 

44 

60 

74 

R 

B 

B 

/r 

R 

9 

25 

41 

57 

74 

R 

5 

2/ 

37 

53 

74 

R 

/ 

17 

33 

49 

74 

R 

13 

29 

45 

6/ 

74 

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3 

B 

/r 

R 

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26 

42 

58 

K 

R 

6 

ZZ 

38 

54 

n 

R 

2 

is 

34 

50 

74 

R 

14 

30 

46 

62 

74 

R 

3 

B 

K 

R 

n 

27 

43 

59 

K 

R 

7 

23 

39 

55 

74 

R 

3 

19 

35 

5/ 

K 

R 

15 

3/ 

47 

63 

K 

R 

B 

B 

74 

R 

12 

28 

44 

60 

K 

R 

8 

24 

40 

56 

n 

R 

4 

20 

36 

52 

74 

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16 

32 

48 

64 

74 

R 

3 

B 

74 

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/3 

29 

45 

61 

74 

R 

9 

25 

41 

57 

74 

R 

5 

2! 

37 

53 

74 

R 

/ 

77 

33 

49 

K 

/? 

B 

B 

A 

R 

M- 

30 

46 

62 

74 

R 

10 

26 

42 

58 

74 

R 

6 

21 

38 

54 

74 

R 

2 

18 

34 

so 

74 

R 

3 

B 

74 

R 

15 

3/ 

47 

63 

74 

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// 

27 

43 

59 

74 

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7 

23 

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55 

74 

R 

3 

19 

35 

5/ 

74 

74 

B 

B 

74 

R 

16 

32 

48 

64 

R 

R 

12 

22 

41 

60 

K 

R 

8 

Z4 

40 

56 

K 

R 

4 

20 

36 

52 

74 

R 

B 

B 

74 

R 

65 

73 

8J 

89 

K 

R 

77 

73 

87 

95 

K 

R 

69 

77 

85 

93 

74 

R 

67 

75 

83 

91 

K 

R 

B 

B 

n 

R 

66 

74 

82 

90 

H 

R 

72 

80 

38 

96 

K 

R 

70 

76 

86 

94 

n 

R 

68 

76 

84 

92 

n 

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B 

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74 

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67 

75 

83 

SI 

K 

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65 

73 

81 

89 

74 

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7/ 

79 

87 

95 

14 

R 

69 

77 

85 

93 

n 

R 

B 

B 

74 

R 

68 

76 

84 

32 

74 

R 

66 

14 

82 

30 

R 

R 

72 

80 

88 

96 

K 

R 

70 

IQ 

86 

94 

74 

R 

B 

B 

n 

R 

69 

77 

85 

93 

74 

R 

67 

75 

83 

9! 

n 

n 

65 

73 

81 

89 

K 

R 

7/ 

79 

67 

95 

n 

R 

3 

B 

74 

R 

70 

78 

86 

34 

K 

R 

68 

76 

84 

92 

n 

R 

66 

74 

82 

90 

74 

R 

72 

80 

68 

96 

K 

R 

B 

B 

K 

R 

7/ 

79 

87 

95 

ff 

R 

69 

77 

85 

93 

n 

R 

67 

75 

83 

5/ 

n 

R 

65 

73 

81 

89 

n 

R 

B 

B 

74 

R 

72 

80 

88 

96 

H 

R 

70 

78 

66 

94 

n 

R 

68 

76 

84 

32 

74 

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66 

74 

82 

90 

71 

R 

3 

Figure  5. — Planting  Plan  of  Oats  Variety  and  Strain  Tests  1921. 
Legend : B,  border.  K,  Kherson  check.  R,  Red  Rustproof  check.  Numbers 
1-64,  planting  numbers  of  oats  strains,  as  given  in  Table  8.  Numbers  65-96, 
planting  numbers  of  oats  varieties,  as  given  in  Table  7. 


Experiments  in  Field  Plot  Technic 


21 


Table  7 —Yields  oe  Oats  Varieties. 
In  Bushels  per  Acre.  1921. 


Planting 

number 

Variety 

Average  yield 
3 Interior  Rows  5 Rows 

65 

Burt 

49.13 

51.94 

66 

Canadian 

25.31 

25.13 

67 

C.  I.  603 

22.50 

23.06 

68 

Culberson 

24.75 

25.13 

69 

Danish  Island 

19.69 

19.13 

70 

Early  Dakota 

21.56 

21.56 

71 

Early  Gothland 

23.44 

22.13 

72 

Garton  748 

21.00 

20.81 

73 

Green  Russian 

26.06 

26.25 

74 

Irish  Victor 

29.81 

32.06 

75 

Joanette 

19.31 

19.69 

76 

Fulghum  042 

45.19 

47.44 

77 

Monarch 

29.63 

31.88 

78 

Monarch  Selection 

35.63 

36.38 

79 

Scottish  Chief 

26.63 

27.38 

80 

Silvermine  050 

31.69 

32.06 

81 

Silvermine  Selection 

22.13 

24.94 

82 

Sparrowbill  (C) 

15.38 

14.63 

83 

Sterilis  Selection 

38.63 

36.94 

84 

Storm  King 

20.06 

17.63 

85 

Swedish  Select  057 

21.00 

19.50 

86 

Fulghum  065 

42.00 

44.81 

87 

Fulghum  0113 

42.00 

45.38 

88 

Silvermine  0115 

25.13 

24.94 

89 

Silvermine  0117 

21.75 

22.69 

90 

Fulghum  0124 

45.38 

48.38 

91 

Fulghum  0145 

39.19 

41.81 

92 

Fulghum  0149 

42.75 

47.06 

93 

Fulghum  0151 

39.75 

43.88 

94 

Fulghum  0152 

39.75 

42.38 

95 

Silvermine  0165 

28.31 

26.81 

96 

Swedish  Select  0165 

20.81 

18.56 

Mean 

29.85 

30.70 

22 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


Table  8. — Yields  of  Oats  Strains  (Red  Rustproof  and  Kherson). 
In  Bushels  per  Acre.  1921. 


Red  Rustproof  strains 

Average  yields 
Planting  3 Interior  5 

number  Strain  Rows  Rows 


1 

066 

24.00 

23.25 

3 

067 

24.00 

21.75 

5 

068 

23.25 

23.44 

7 

069 

19.31 

18.00 

9 

072 

18.38 

18.75 

11 

074 

22.31 

20.63 

13 

075 

24.19 

22.13 

15 

0118 

16.50 

16.31 

18 

0119 

22.31 

21.38 

20 

0120 

21.19 

19.69 

22 

0122 

19.13 

17.81 

24 

0125 

21.00 

19.88 

26 

0126 

25.31 

22.50 

28 

0128 

20.44 

20.25 

30 

0129 

21.94 

21.56 

32 

0130 

21.75 

20.25 

33 

0131 

24.56 

23.25 

35 

0132 

17.63 

19.13 

37 

0133 

18.94 

18.75 

39 

0134 

16.50 

15.75 

41 

0135 

17.63 

15.65 

43 

0136* 

32.44 

33.19 

45 

0141 

21.94 

21.19 

47 

0163 

13.88 

12.94 

50 

0169 

15.38 

14.44 

52 

0181 

19.88 

18.00 

54 

0182 

19.88 

19.13 

56 

0183* 

41.44 

43.31 

58 

0383 

23.63 

24.00 

60 

0391 

29.44 

30.19 

62 

0394 

22.31 

21.56 

64 

0395 

23.25 

21.94 

Mean 

21.00 

20.12 

♦Not  taxonomically  Red  Rustproof.  Ex- 
cluded from  average. 


Kherson  strains 

Average  yields 

Planting  3 Interior  5 


Number  Strain 

Rows 

Rows 

2 

023 

35.25 

36.38 

4 

040 

36.57 

37.50 

6 

041 

36.56 

38.81 

8 

052 

38.06 

38.81 

10 

053 

39.75 

42.00 

12 

079 

32.63 

34.88 

14 

080 

35.44 

38.25 

16 

082 

40.88 

41.44 

17 

083 

35.44 

38.25 

19 

085 

38.25 

41.81 

21 

086 

36.75 

37.69 

23 

Mixture** 

33.75 

36.38 

25 

088*** 

27.00 

27.56 

27 

089 

30.94 

31.69 

29 

090 

36.38 

38.06 

31 

091 

30.19 

31.88 

34 

094 

31.69 

33.56 

36 

095 

38.81 

39.75 

38 

096 

36.38 

38.06 

40 

097 

31.31 

32.25 

42 

098 

38.63 

39.19 

44 

099 

38.81 

38.25 

46 

0100 

40.13 

42.75 

48 

0155 

37.50 

38.63 

49 

0157 

43.69 

45.00 

51 

0158 

34.69 

35.25 

53 

0159 

33.38 

33.94 

55 

0160 

30.19 

31.13 

57 

0161 

34.69 

36.00 

59 

0162 

25.31 

25.69 

61 

0167 

40.69 

39.94 

63 

0174 

36.75 

38.25 

Mean 

35.79 

37.14 

♦♦Mixture  of  strains 

082,  094, 

0100, 

0174. 

♦♦♦Not 

taxonomically 

Kherson. 

Ex- 

eluded  from  average. 


Experiments  in  Field  Plot  Technic 


23 


COMPETITION  AS  A SOURCE  OF  ERROR  IN  PRELIMINARY 

TESTS. 

Previous  Investigation. — The  possibility  of  error  from  competi- 
tion in  single-row  tests  was  noted  by  Montgomery14  in  1913,  in  the 
following  passage : 

“In  1908  it  was  observed  that  a certain  strain  of  early  wheat  in  a series 
of  row  plats  made  a very  poor  appearance  at  harvest  time,  while  the  same 
strain  planted  in  centgeners  made  a much  better  comparative  showing.  Ap- 
parently the  larger  and  faster  growing  strains  on  each  side,  the  rows  being 
only  8 inches  apart,  exercised  some  competitive  effect.  This  effect  of  com- 
petition has  been  noted  for  two  years  since.  Also  in  certain  variety  tests 
of  oats,  grown  in  row  plats  10  inches  apart,  the  same  effect  was  noted. 
Exact  data  cannot  be  given  on  this  point,  as  the  results  from  the  series  of 
plats  planted  in  1909  and  in  1910  for  this  purpose  were  seriously  impaired 
by  unfavorable  conditions;  but  Table  XVIII,  giving  results  from  adjacent 
row  plats  sown  at  different  rates,  shows  that  the  800-seed  rate  made  a marked 
increase  over  the  700-seed  rate,  while  in  a similar  series  of  blocks  (Table 
XIX),  sown  at  the  same  rate,  this  marked  increase  was  not  noted.  Since  the 
800-seed  row  was  always  adjacent  to  the  400-seed  row,  it  may  have  had  some 
advantage  on  this  account.  Danger  from  this  source  can  probably  be  avoided 
if  care  is  taken  to  plant  only  similar  varieties  in  adjacent  rows.  Where  the 
block  plat  is  used  this  source  of  error  is  eliminated.” 

Hayes  & Arny4  found  considerable  competition  between  rod-rows 
grown  one  foot  apart.  Three-row  plots  were  used  in  variety  tests  of 
winter  wheat,  spring  wheat,  barley,  and  oats,  and  the  yields  of  each 
row  determined  separately,  in  1916.  The  comparative  yield  of  the 
border  rows  in  each  plot  was  then  correlated  with  the  comparative 
height  and  yield  of  the  adjacent  rows.  There  was  some  effect  on  the 
yield  of  border  rows  due  to  the  height  of  adjacent  rows  in  the  case 
of  barley  and  winter  wheat.  The  results  were  variable  in  different 
plots.  In  the  case  of  oats  the  effect  of  height  was  rather  obscure,  and 
in  the  case  of  spring  wheat  it  was  not  apparent.  The  yield  of  adjacent 
rows  appeared  to  be  of  some  importance  in  the  barley  tests  and  in 
some  of  the  spring  wheat  tests.  These  results  led  to  the  adoption  of 
3-row  plots  with  discarded  border  rows  for  preliminary  testing  at 
the  Minnesota  Station. 

Love  and  Craig*  in  describing  the  methods  used  in  cereal  investiga- 
tions at  the  Cornell  Station  describe  the  single-row  test  and  add : “In 
order  to  prevent  any  effect  which  may  be  caused  by  two  unlike  sorts 
growing  together  the  different  strains  are  arranged  according  to 
earliness  and  other  characters  so  as  to  reduce  this  source  of  error  to  a 
minimum.” 


24 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


Kiesselbach5’ 7 has  published  rather  extensive  data  on  the  compe- 
tition between  adjacent  rod-rows.  In  his  experiments  the  crops  were 
compared  in  alternating  single-row  plots  and  in  alternating  5-row 
blocks,  each  replicated  fifty  times.  In  some  cases  the  border  rows  of 
5-row  plots  were  discarded.  The  deviation  of  the  result  in  the  test 
in  single-row  plots  from  that  of  the  test  in  5-row  plots  is  regarded  as  the 
measure  of  the  effect  of  competition.  The  comparative  yields  of  va- 
rieties of  wheat  and  oats  in  alternating  single  rows  and  in  alternating 
5-row  plots  are  shown  in  Table  9,  from  Kiesselbach7. 


Tabu;  9. — Relative  Yields  of  Two  Small  Grain  Varieties  When  Compared 
in  Alternating  Rows  and  in  Alternating  5-Row  Plats  (Kiesselbach). 


Wheat 

Oats 

Average  yield  of  l 

50  plats 

Average  yield  of  50  plats 

Year  and 

Alternating 

Alternating 

Year  and 

Alternating 

Alternating 

variety 

single  rows 

5-row  blocks* 

Variety 

single  rows 

5-row  blocks’1 

% 

% 

% 

% 

1913 

1913 

Turkey 

100 

100 

Kherson 

100 

100 

Big  Frame 

107 

97 

Burt 

130 

112 

1914 

1914 

Turkey 

100 

100 

Kherson 

100 

100 

Big  Frame 

85 

97 

Burt 

139 

101 

1913 

1913 

Turkey 

100 

100 

Kherson 

100 

100 

Neb.  No. 

107 

107 

Swedish 

82 

77 

28 

Select 

1914 

1914 

Turkey 

100 

100 

Kherson 

100 

100 

Neb.  No. 

63 

85 

Swedish 

89 

93 

28 

Select 

♦Yield  based  on  3 inner  rows  of  5-row  plats  in  1914. 

Kiesselbach  also  submits  interesting  data  on  the  competition  of 
pure  line  selections  of  the  same  variety.  It  might  be  supposed  that 
such  strains,  being  similar  in  varietal  characteristics,  would  be  little 
affected  by  competition,  and  could  therefore  be  safely  compared  in 
single-row  plots.  The  average  relative  yields  of  three  strains  of  Turkey 
wheat  in  single  rows  and  in  blocks  for  two  seasons,  however,  showed 
that  the  two  better  strains  were  favored  approximately  20  per  cent  and 
15  per  cent,  respectively,  at  the  expense  of  the  poorer  strain,  in  the 
single-row  test.  A strain  which  yielded  26  per  cent  more  than  an- 
other in  the  single-row  test  yielded  only  6 per  cent  more  in  the  block 


Experiments  in  Field  Plot  Technic 


25 


test.  Kiesselbach  has  therefore  adopted  the  practice  of  testing  such 
strains  in  5-row  blocks  replicated  ten  times  instead  of  in  single-row 
plots. 

Love8  has  criticized  these  results  because  in  some  cases  at  least 
the  rows  ran  east  and  west  rather  than  north  and  south.  He  states 
that  in  experiments  at  Ithaca,  New  York,  there  is  little  competition  be- 
tween varieties  grown  in  single  rows,  when  the  rows  run  north  and 
south.  “In  order  to  obviate  any  criticism  of  this  method,”  he  adds,  “it 
might  be  well  to  follow  the  plan  of  arranging  varieties  so  that  late 
sorts  are  grown  together  and  the  earlier  ones  together.  In  other  words, 
the  different  sorts  could  be  so  arranged  that  they  grade  into  one  another 
as  regards  yield,  earliness,  and  the  like.”  To  this  Kiesselbach8  replies 
that  in  some  of  his  competition  studies  the  rows  ran  north  and  south 
and  in  others  east  and  west,  and  that  striking  competition  occurred  in 
both  cases.  He  adds  that  although  error  resulting  from  row  compe- 
tition would  undoubtedly  be  reduced  by  grouping  varieties  of  sim- 
ilar growth  habits  together,  it  appears  that  varieties  fairly  similar 
in  growth  habit  may  vary  for  some  reason  in  relative  competitive  qual- 
ity. 

Experimental  Results. — Some  further  evidence  on  competition 
as  a source  of  error  in  plot  experiments  is  afforded  by  a study  of  the 
relative  yields  of  border  rows  and  interior  rows  in  the  5-row  blocks  used 
in  these  preliminary  tests.  It  should  be  remembered,  of  course,  that  the 
effect  on  yield  would  be  decidedly  greater  in  single  rows  exposed  to  com- 
petition on  both  sides  than  in  these  border  rows,  which  compete  with 
another  sort  on  only  one  side.  The  extent  of  the  error  from  compe- 
tition in  such  border  rows  is  of  interest  in  determining  whether  it  is 
necessary  to  discard  the  border  rows  of  small  blocks.  When  5-row 
blocks  are  used,  even  if  the  border  rows  are  not  discarded,  the  relative 
effect  of  competition  is  greatly  reduced,  since  only  two  of  the  five 
rows  are  subject  to  varietal  competition  and  these  are  exposed  only 
on  one  side.  If  this  results  in  reducing  the  error  from  competition  to 
a low  point,  or  if  varieties  can  be  so  arranged  as  to  give  this  result, 
it  may  be  advisable  in  practice  to  harvest  5-row  blocks  entire,  thus 
avoiding  the  principal  objection  to  the  use  of  border  rows — the  loss  of 
a considerable  portion  of  the  experimental  area. 

Competition  is  particularly  important  as  a source  of  error  because 
of  the  fact  that  it  tends  to  affect  replicate  plots  similarly,  and  conse- 
quently does  not  necessarily  increase  plot  variability.  For  this  reason 
it  is  likely  to  escape  detection,  and,  when  it  is  involved  in  an  experi- 
ment, its  effect  cannot  be  measured.  There  is  no  'great  objection  to  a 
considerable  experimental  error  from  plot  variability  in  field  experi- 


26 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


ments,  if  the  experimenter  determines  the  extent  of  the  error  and  draws 
his  conclusions  accordingly.  But  a preliminary  variety  test  in  which 
error  from  competition  is  not  controlled  may  be  very  nearly  worthless 
as  an  indication  of  the  relative  value  of  varieties  for  field  conditions, 
because  actually  the  relative  values  of  the  varieties  tested  may  frequent- 
ly differ  by  50  or  100  per  cent  from  the  values  determined  in 
the  test,  without  the  slightest  indication  in  the  experimental  results. 

Illustrations  of  Effects  of  Competition. — The  error  from  competi- 
tion may  be  illustrated  by  numerous  examples  from  each  of  the  eight 
tests  here  reported.  An  extreme  case  is  the  effect  of  competition  on 
the  relative  yield  of  wheat  and  rye.  Two  varieties  of  rye,  common 
rye  and  Rosen  rye,  were  included  in  the  wheat  variety  test,  for  com- 
parison with  wheat.  The  average  yields  of  Rosen  rye  and  of  the  va- 
rieties of  wheat  adjoining  it  on  either  side,  in  interior  rows  and  com- 
peting border  rows  of  the  four  series,  were  as  follows: 


Season  Variety 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

i 

[Niagara  (Wheat) 

13.8 

67 

10.0 

33 

1 

f30.4 

100 

1920< 

[ Rosen  (Rye) 

20.7 

100 

\ 

— 

[27.7 

100 

Velvet  Chaff  No.  2 (Wheat 

14.1 

68 

9.8 

35 

1 

[Red  Hussar  (Wheat) 

14.3 

80 

11.6 

59 

1 

[19.7 

100 

1921-j 

Rosen  (Rye) 

17.9 

100 

\ 

— 

[25.4 

100 

Poole  (ck)  (Wheat) 

1 

11,8 

66 

11.2 

44 

The  disturbance  of  the  true  comparative  value  of  the  varieties 
by  competition  may  be  determined  by  comparing  their  relative  yields 
in  interior  rows  and  in  border  rows.  Thus  Niagara  wheat  in  1920 
yielded  67  per  cent  as  much  as  Rosen  rye  in  plots  protected  from  com- 
petition, but  only  33  per  cent  as  much  in  rows  not  protected  from 
competition.  Similarly  the  yield  of  Velvet  Chaff  No.  2 wheat  was  re- 
duced from  68  per  cent  to  35  per  cent  by  competition  with  Rosen  rye. 
In  the  following  season  the  reduction  in  yield  of  the  two  varieties  of 
wheat  adjoining  Rosen  rye  (Red  Hussar  and  Poole)  was  not  so 
great,  but  was  still  decidedly  significant.  This  clear  case  of  compe- 


Experiments  in  Eield  Plot  Technic 


27 


tition  serves  to  illustrate  the  phenomenen,  although  the  competition 
between  wheat  and  rye  has  little  significance  in  itself  as  regards  va- 
riety tests  in  general,  since  wheat  and  rye  are  not  commonly  included 
in  the  same  test. 

Ordinarily  the  competition  between  varieties  of  the  same  crop 
is  not  so  extreme.  There  are,  however,  a number  of  cases  in  which 
a variety  of  wheat  or  oats  profited  almost  as  extremely  in  competition 
with  other  varieties  of  the  same  crop  as  did  the  rye  in  competition 
with  wheat  in  the  cases  cited  above.  The  wheat  variety,  Michigan 
Wonder  No.  116,  which  grew  between  two  other  wheat  varieties, 
Leap's  Prolific  and  Poole  Selection,  in  1921,  gave  the  following  results, 
as  an  average  of  the  four  series: 


Variety 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

Leap’s  Prolific 

14.9 

91 

9.9 

53 

f 18.8 

100 

Michigan  Wonder  No.  116 

16.4 

100 

\ 

— 

1 

[21.7 

100 

Poole  Selection 

15.3 

1 

93 

j 

11.5 

53 

The  effect  of  competition  in  this  case  is  almost  as  pronounced 
as  in  the  case  of  the  rye,  although  the  three  wheat  varieties  concerned, 
when  protected  from  competition,  gave  almost  equal  yields  and  differed 
little  in  date  of  heading,  date  of  maturity,  and  height.  In  this  case 
a small  difference  in  actual  value  between  the  varieties,  as  indicated 
by  their  yields  when  protected  from  competition,  is  greatly  increased 
when  their  yields  in  adjacent  single  rows  are  compared. 

A striking  case  of  competition  in  the  oats  variety  test  of  1921  was 
that  of  the  three  varieties  Sterilis  Selection,  Fulghum,  and  Kherson, 
the  check  variety.  Their  average  yields  were  as  follows : 


Variety 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

Sterilis  Selection 

38.63 

99 

28.50 

58 

f 48.75 

100 

Fulghum 

39.19 

100 

\ 

— 

[ 42.94 

100 

Kherson  (check) 

40.69 

104 

34.18 

70 

28 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


These  three  varieties,  which  gave  almost  equal  yields  in  rows 
protected  from  competition,  differed  decidedly  in  their  yields  in  ad- 
jacent rows.  Although  Kherson  outyielded  Fulghum  4 per  cent  in 
plots  protected  from  competition,  its  yield  was  30  per  cent  less  than 
that  of  Fulghum  in  single  rows  not  protected  from  competition. 

Extreme  effects  of  competition  were  shown  in  very  numerous 
cases  in  the  tests  of  Kherson  and  Red  Rustproof  strains  in  1921.  An 
example  from  this  plot  is  the  following: 


Strain 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

0169  (Red  Rustproof  check) 

18.38 

77 

22.31 

119 

1 

[18.75 

100 

067  (Red  Rustproof) 

24.00 

100 

! 

1 

— 

: 1 

[17.44 

100 

085  (Kherson) 

38.25 

159 

i 

51.94 

298 

The  extreme  advantage  of  the  Kherson  strain  in  competition  with 
the  Red  Rustproof,  increasing  its  margin  of  superiority  from  59  per 
cent  to  198  per  cent,  is  particularly  striking.  Probably  even  more 
significant  is  the  effect  of  competition  between  the  two  Red  Rustproof 
strains,  resulting  in  the  conversion  of  a 23  per  cent  loss  to  a 19  per 
cent  gain. 

All  of  the  cases  cited  above  are  taken  from  plots  in  which  the 
rows  ran  east  and  west.  Some  examples  of  varietal  competition  from 
tests  in  rows  running  north  and  south  are  the  following: 

In  the  barley  variety  test,  the  variety  Featherston  1118  occurred 
between  Red  River  973  and  Oderbrucker  (the  check  variety).  The 
average  yields  of  these  three  varieties  in  the  three  series  were  as  follows : 


Variety 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

Red  River  973 

27.25 

96 

31.20 

127 

f24.60 

100 

Featherston  1118 

28.25 

100 

j 

— 

(25.65 

100 

Oderbrucker  (ck) 

34.87 

123 

42.19 

164 

Experiments  in  Field  Plot  Technic 


29 


In  this  case  the  advantage  of  Oderbrucker  over  Featherston  was 
almost  tripled  by  competition,  and  Red  River,  which  yielded  less  than 
Featherston  in  the  interior  rows,  excelled  it  materially  in  yield  in  the 
border  rows. 

The  oats  varieties  tested  in  rows  running  north  and  south  in  1919 
showed  marked  effects  of  competition  in  several  cases.  The  follow- 
ing will  serve  as  an  example: 


Variety 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

Kherson  Selection 

61.3 

111 

84.7 

[49.4 

171 

100 

Fulghum  042 

57.1 

100 

[50.7 

100 

Lincoln 

50.3 

88 

57.5 

113 

In  this  case  Lincoln,  yielding  12  per  cent  less  than  Fulghum  in  in- 
terior rows,  yielded  13  per  cent  more  than  Fulghum  in  border  rows; 
while  the  advantage  of  Kherson  Selection  over  Fulghum  was  in- 
creased from  11  per  cent  to  71  per  cent. 

Marked  competition  is  hardly  to  be  expected  in  the  oats  strain  test 
of  1919,  regardless  of  the  direction  of  the  rows,  because  of  the  sim- 
ilarity of  the  strains  in  varietal  characters.  Three  strains  which  proved 
to  be  taxonomically  unlike  Red  Rustproof  were  included  in  this  test, 
and  each  of  these  shows  clearly  the  effects  of  competition.  For  ex- 
ample, strain  0124,  which  was  classified  as  Fulghum,  gave  the  follow- 
ing yields  in  comparison  with  the  adjoining  strains,  0127,  classified  as 
Kherson,  and  0133,  classified  as  Red  Rustproof : 


Strain 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

0127  (Kherson) 

38.55 

60 

32.40 

48 

f66.83 

100 

0124  (Fulghum) 

C3.90 

300 

— 

[78.75 

100 

0133  (Red  Rustproof) 

48.00 

75 

43.20 

55 

30 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


Moreover,  the  Red  Rustproof  strains  showed  competitive  effects 
among  themselves  to  some  extent,  though  not  so  conspicuously  as  dif- 
ferent varieties.  For  example  the  strains  0122  and  0123,  which  were 
taxonomically  identical,  yielded  as  follows: 


Strain 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

0122  (Red  Rustproof) 

0123  (Red  Rustproof) 

47.85 

53.55 

89 

10O 

1 

55.13 

49.28 

112 

100 

Strain  0122  which  was  apparently  11  per  cent  inferior  to  strain 
0123  in  the  yields  of  interior  rows,  appeared  to  be  12  per  cent  superior 
to  the  same  strain  in  the  yields  of  their  adjacent  border  rows. 

In  the  wheat  mixture  test  of  1920  also  the  rows  ran  north  and 
south.  An  example  of  competition  from  this  test  is  the  following: 


Variety 

Yield  in 
interior  rows 

Yield  in  competing 
border  rows 

Bushels 

Relative 

Bushels 

Relative 

Poole  (check) 

15.1 

81 

14.5 

60 

f24.2 

100 

Michigan  Wonder  No.  221 

18.6 

100 

\ 

— 

[21.4 

100 

Kanred 

13.6 

73 

1 

12.5 

58 

In  this  case  also  differences  in  yield  were  increased  by  compe- 
tition. 

The  individual  cases  cited  above  will  serve  to  show  the  existence 
of  competition  as  a source  of  error  in  these  tests.  As  a result  of 
competition  the  differences  between  varieties  may  be  increased  or  de- 
creased, and  in  some  cases  a material  advantage  in  yield  may  be  con- 
verted to  a material  disadvantage.  The  phenomenon  occurs,  under 
conditions  at  Columbia,  whether  the  rows  run  north  and  south  or 
east  and  west.  Of  course  it  is  not  true  that  all  of  the  difference  in 
yield  between  border  rows  and  interior  rows  is  necessarily  caused 
by  varietal  competition.  Some  variation  in  the  yield  of  adjacent  rod- 
rows  will  occur  regardless  of  competition.  When  the  means  of  only 


Experiments  in  Field  Plot  Technic 


31 


four  determinations  are  compared  the  effect  of  this  variability  may  be 
considerable.  If  a field  uniformly  seeded  to  a single  strain  were  har- 
vested in  rod-rows  and  assumed  to  be  made  up  of  several  different 
varieties  each  in  four  distributed  plots,  doubtless  the  average  border 
yield  would  differ  materially  from  the  average  interior  yield  in  several 
“varieties.”  It  is  not  however,  likely,  that  such  differences  as  those 
cited  above  would  be  caused  by  chance  variability.  Nevertheless,  no 
final  conclusions  regarding  competition  as  a source  of  error  should  be 
drawn  from  such  individual  cases.  The  extent  of  error  from  compe- 
tition is  better  shown  in  the  average  differences  between  border  yields 
and  interior  yields,  and  in  the  mean  coefficients  of  competition  for 
complete  tests.  They  are  given  in  the  next  section. 

Relation  of  Competition  to  Various  Characteristics  of  the  Com- 
peting Varieties. — It  is  essential  that  competition  be  eliminated  by  the 
use  of  border  rows,  or  counteracted  by  some  such  means  as  grouping 
varieties.  The  latter  is  decidedly  the  preferable  method,  from  the 
standpoint  of  economy,  if  satisfactory  results  may  be  obtained  by  its 
use.  But  competition  cannot  be  effectively  controlled  by  grouping 
varieties  unless  there  is  a close  correlation  between  competitive  value 
and  some  character  like  earliness  or  height,  which  may  be  known  in 
advance.  Determinations  of  the  correlation  between  competitive  ef- 
fects and  various  characteristics  of  the  varieties  have  therefore  been 
made  for  each  of  the  tests.  The  preliminary  determinations  were  made 
as  follows : 

(1)  The  average  yield  in  interior  rows  and  the  average  yield 
in  the  border  rows  on  each  side  for  all  replicate  plots  of  each  variety 
or  strain  was  determined.  The  replicate  plots  thus  averaged  were 
grown  between  the  same  varieties  in  each  series,  and  it  may  be  assumed 
therefore  that  their  border  rows  were  subject  to  the  same  competition. 
In  the  following  discussion  of  competition  each  individual  case  repre- 
sents the  mean  of  all  the  replicate  plots  of  the  test  in  question.  For 
example,  when  it  is  stated  that  the  correlation  between  competition 
and  yield  is  determined  in  a test  in  which  one  hundred  cases  of  compe- 
tition are  involved,  each  of  the  hundred  cases  represents  the  mean  of 
three  or  four  determinations  in  replicate  plots.  In  most  cases  the 
number  of  replicate  plots  was  four.  In  the  barley  test  of  1919  only 
three  series  were  grown,  and  in  the  oats  variety  test  of  1919,  though 
four  series  were  grown,  only  three  could  be  used  because  one  border 
row  of  each  variety  in  the  first  series  was  harvested  for  seed  and 
laboratory  material. 

(2)  Corresponding  average  yields  were  determined  for  check 
plots,  those  adjoining  the  same  variety  being  averaged  together.  For 


32 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


example,  in  the  wheat  variety  test  diagrammed  in  figure  1 the  four 
check  plots  which  adjoined  variety  1 (one  in  each  series)  were  aver- 
aged together,  the  four  adjoining  variety  2,  the  four  adjoining  variety 
3,  etc.  The  four  check  plots  adjoining  varieties  89,  90,  91,  etc.  were 
similarly  averaged. 

(3)  The  average  yield  of  each  border  row  for  each  variety  was 
converted  to  the  percentage  of  the  average  yield  of  the  same  variety 
in  its  interior  rows.  These  yields  of  border  rows  in  percentage  will 
be  referred  to  as  “relative  border  yields.”  The  relative  border  yield 
gives  a rough  indication  of  the  effect  of  competition  on  the  variety. 
When  it  is  above  100,  the  variety  yielded  more  in  border  rows  (subject 
to  competition)  that  in  interior  rows  (protected  from  competition). 
When  it  is  below  100,  the  border  yield  was  less  than  the  interior  yield, 
in  proportion. 

(4)  An  approximate  measure  of  the  competition  between  each 
pair  of  adjacent  varieties  was  obtained  by  dividing  the  higher  relative 
border  yield  by  the  lower,  in  the  case  of  their  adjacent  border  rows, 
and  substracting  100  from  the  result.  When  the  variety  on  the  left 
has  a higher  relative  border  yield,  this  is  given  a positive  sign;  in  the 
reverse  case  a negative  sign.  This  figure  is  simply  the  predominance 
of  the  more  strongly  competing  variety  over  the  other  in  percentage  of 
relative  border  yield.  It  will  be  referred  to,  for  convenience,  as  the 
coefficient  of  competition. 

(5)  This  measure  of  competition  was  correlated  with  various 
characteristics  of  the  competing  varieties,  including  the  relative  yields 
in  interior  rows,  the  relative  grain-straw  ratios,  the  relative  dates  of 
heading  and  of  maturity,  and  the  relative  heights.  In  correlating  com- 
petition with  the  relative  yield  of  the  interior  rows,  the  relative  yield 
was  determined  by  dividing  the  higher  yield  by  the  lower,  subtracting 
100,  and  assigning  a positive  or  negative  sign,  as  before.  The  correla- 
tion determined,  therefore,  is  the  correlation  between  the  percentage 
advantage  of  one  variety  over  another  in  competition,  and  the  differ- 
ence in  yield  of  the  two  varieties,  expressed  in  percentage,  when  pro- 
tected from  competition.  Relative  grain-straw  ratios  were  determined 
similarly,  the  ratios  being  first  obtained  by  dividing  the  yield  of  straw 
by  the  yield  of  grain.  Relative  dates  of  heading  and  maturity  and 
relative  heights  were  determined  simply  by  subtracting  the  value  for 
one  variety  from  the  value  for  the  other.  In  each  case,  of  course,  the 
sign  was  determined  in  the  same  way. 

A simple  example  explained  in  detail  may  serve  to  make  this 
method  clear.  In  the  wheat  variety  test  of  1921  the  varieties  Fultz 
(Bayer),  Michigan  Amber,  and  Michigan  Wonder  No.  211  occurred 


Experiments  in  Field  Plot  Technic 


33 


in  the  order  named  in  four  distributed  sections  of  the  field.  The  aver- 
age yields  of  these  varieties  in  the  four  series,  in  bushels  per  acre,  for 
border  rows  and  for  interior  rows,  are  shown  below,  together  with 
the  average  dates  of  heading,  dates  of  maturity,  and  heights,  also  de- 
termined for  the  four  series. 


23.  Fultz  (Bayer) 

39.  Michigan  Amber 

55.  Michigan  Wonder 
No.  211 

Row 

1 

Row 
2,  3,  4 

Row 

5 

Row 

1 

Row 
2,  3,  4 

Row 

5 

Row 

1 

Row 
2,  3,  4 

Row 

5 

Average 

yields 

10.8 

bu. 

12.2 

bu. 

13.1 

bu. 

13.3 

bu. 

14.9 

bu. 

14.5 

bu. 

19.8 

bu. 

18.1 

bu. 

19.4 

bu. 

Average 
date  of 
heading 

21* 

21* 

19* 

Average 
date  of 
maturity 

47* 

48* 

47* 

Average 

height 

43  in. 

42  in. 

43  in. 

* Dates  of  heading  and  maturity  are  the  numbers  of  days  after  April  30.  Thus  1 is 
May  1,  32  is  June  1,  47  is  June  16,  etc. 


Now  dividing  the  yields  in  border  rows  by  the  yields  of  the  same 
varieties  in  interior  rows,  we  obtain  the  relative  border  yields,  which 
are  substituted  in  the  table  below  for  the  border  yields  in  bushels. 
To  determine  the  degree  of  competition  between  the  varieties  Fultz  and 
Michigan  Amber  we  divide  the  larger  relative  border  yield  (107)  by  the 
smaller  (89)  and  subtract  100,  giving  20  per  cent.  Since  in  this  case  the 
relative  border  yield  of  the  variety  on  the  left  is  higher,  the  difference  is 
given  a minus  sign.  Similarly  a value  of  +12  per  cent  is  obtained  for 
the  competition  between  Michigan  Amber  and  Michigan  Wonder  No. 
211.  These  figures  mean  that  the  relative  border  yield  of  Fultz  ex- 
ceeded that  of  Michigan  Amber  by  20  per  cent  in  their  competing 
border  rows,  while  that  of  Michigan  Wonder  exceeded  that  of  Michi- 
gan Amber  by  12  per  cent. 

The  relative  yields  of  these  varieties  are  obtained  similarly, — 
in  the  first  case  by  dividing  14.9  by  12.2  (+22%)  and  in  the  second 
case  by  dividing  18.1  by  14.9  (+21%).  Both  values  are  positive  be- 
cause in  each  case  the  yield  of  the  variety  on  the  left  is  higher  than 
that  of  the  variety  on  the  right.  The  difference  in  dates  of  heading, 
maturity,  and  height  are  obtained  simply  by  subtraction,  being  positive 
when  the  value  of  the  variety  on  the  right  is  greater  and  negative  when 


34 


Missouri  Agr.  Exp.  Sta.  Research  Bueletin  49 


the  value  of  the  variety  on  the  left  is  greater.  The  figures  ready  for 
correlation  study  will  then  appear  as  follows : 


23.  Fultz 
(Bayer) 

Row  Row  Row 
1 2,3,4  5 

Compe- 

tition 

data 

39.  Michigan 
Amber 

Row  Row  Row 

1 2,  3,  4 5 

Compe- 

tition 

data 

5 5.  Michigan  Wonder 
No.  211 

Row  Row  Row 

1 2,  3, 4 5 

Average 

yield 

89 

12.2 

107 

-20% 

+22% 

89 

14.9 

97 

+ 12% 
+21% 

109 

18.1 

107 

Average 
date  of 
heading 

21 

0 

21 

—2 

19 

Average 
date  of 
maturity 

47 

+ 1 

48 

0 

47 

Average 

height 

43 

— 1 

42 

+ 1 

43 

The  columns  headed  “competition  data”  show  the  relation  of  the 
effect  of  competition  to  the  yield,  earliness,  and  height  of  the  competing 
varieties.  For  example,  Michigan  Amber  was  at  a disadvantage  of  20 
per  cent  in  competition  with  Fultz,  though  it  was  22  per  cent  superior 
in  yield  when  protected  from  competition.  It  headed  the  same  day, 
matured  one  day  later,  and  was  one  inch  shorter.  After  correspond- 
ing data  had  been  prepared  for  all  the  96  varieties  in  this  test,  correla- 
tion tables  with  the  coefficient  of  competition  as  subject  and  relative 
yield,  date  of  heading,  date  of  maturity,  and  height  as  relative  were 
constructed.  Correlations  were  determined  similarly  in  the  other  tests. 
One  of  these  correlation  tables  is  shown  in  figure  6.  In  general,  merely 


o 

o 

o 

o 

o 

o 

o 

o 

o 

o 

o 

CM 

o 

00 

CM 

CM 

to 

00 

© 

7 

1— 1 
1 

1 

1 

1 

| 

rH 

o 

o 

o 

o 

o 

O 

O 

o 

o 

o 

o 

o 

o 

o 

o 

o 

o 

o 

o 

o 

o 

O 

4-» 

/— s 

o 
1— 1 

1 

00 

1 

o 

1 

1 

1 

CM 

tO 

00 

W 

H 

—40 

to 

—60 

1 

1 

—20 

to 

—40 

2 

1 

1 

2 

4 

2 

2 

14 

0 

to 

—20 

1 

1 

7 

10 

5 

2 

26 

0 

to 

20 

1 

2 

6 

10 

8 

3 

5 

35 

20 

to 

40 

2 

3 

4 

11 

2 

1 

1 

24 

40 

to 

60 

1 

1 

3 

5 

60 

to 

80 

1 

1 

1 

3 

Total 

2 

1 

3 

5 

20 

26 

21 

16 

8 

5 

1 

112 

Figure  6.— Correlation  Between  Coefficient  of  Competition  and  Rela- 
tive Yield,  in  Wheat  Variety  Test  1920. 

r=  +.582  ± .043. 


Experiments  in  Field  Plot  Technic 


35 


the  coefficient  of  correlation  and  its  probable  error  are  given,  for  lack 
of  space. 

In  the  barley  variety  test,  1919,  the  effect  of  competition  was  quite 
marked.  The  average  yield  of  border  rows  differed  from  the  average 
yield  of  interior  rows  by  11.13  per  cent,  and  the  mean  coefficient  of 
competition  was  21.30  per  cent.  Attempts  were  made  to  correlate 
competition  with  relative  date  of  heading,  date  of  maturity,  grain-straw 
ratio,  and  yield.  The  correlation  coefficients  determined  are  shown 
in  Table  10,  together  with  the  mean  differences  between  competing 


Table  10. — Correlation  of  Competition  With  Various  Characteristics  in 
Barley  Varety  Test  1919. 


Character 


Date  of  heading 
Date  of  maturity 
Grain-straw  ratio 
Yield 


Mean  difference  be-  Coefficient  of  correlation 

tween  competing  with  competition 

varieties 


4.0  days 
2.6  days 
38.0% 
52.3% 


—.153  ±.120 
—.063  ±.123 
+ .072  ±.122 
+ .442  ±.099 


varieties  in  the  characters  whose  relation  to  competition  was  studied. 

Although  none  of  these  correlations  is  statistically  significant,  in 
the  strictest  sense,  it  is  noticeable  that  the  correlation  between  compe- 
tition and  yield  is  much  greater  than  any  of  the  others,  and  is  equal 
to  about  four  and  one-half  times  its  probable  error.  There  was  ap- 
parently some  tendency  for  the  better  yielding  varieties  to  profit  by 
competition  with  the  poorer  yielders.  On  account  of  the  relatively 
small  number  of  cases  involved  in  this  and  the  other  1919  tests,  the 
probable  errors  are  high,  and  a fairly  high  coefficient  of  correlation 


Table  11. — Correlation  of  Competition  With  Various  Characteristics  in 
Oats  Variety  Test  1919. 


Character 


Date  of  maturity 
Grain-straw  ratio 
Yield 


Mean  difference  be-  Coefficient  of  correlation 

tween  competing  with  competition 

varieties 

3.56  days  —.456  ±.103 

50.2%  —.091  ±.129 

53.5%  +.314  ±.117 


may  consequently  fail  to  attain  statistical  significance.  Such  a co- 
efficient, while  not  establishing  the  correlation,  by  no  means  indicates 
that  the  correlation  does  not  exist. 


36 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


The  oats  variety  test  of  1919  also  showed  distinctly  the  effects 
of  competition.  The  border  rows  in  this  test  differed  in  yield  from 
the  interior  rows  by  1278  per  cent,  on  the  average,  and  the  mean 
coefficient  of  competition  was  27.67  per  cent.  Correlations  were  de- 
termined for  competition  and  relative  yield,  date  of  maturity,  and 
grain-straw  ratio.  Unfortunately  the  dates  of  heading  are  not  avail- 
able for  all  varieties  in  this  test.  The  correlation  coefficients  are  shown 
in  Table  11. 

Again  no  correlations  of  statistical  significance  are  found,  but 
the  relation  of  yield  and  earliness  of  maturity  to  competing  strength 
is  at  least  suggestive.  There  was  a tendency  for  early  and  high-yielding 
varieties  to  profit  by  competition  at  the  expense  of  later  and  lower-yield- 
ing varieties,  but  the  number  of  varieties  was  too  small  to  permit  the 
drawing  of  positive  conclusions. 

The  oats  strains  grown  on  the  same  field  showed  much  less  strik- 
ingly the  effects  of  competition.  The  mean  difference  in  yield  be- 
tween border  rows  and  interior  rows  in  these  15  strains  was  only  6.50 
per  cent  and  the  mean  coefficient  of  competition  only  13.11  per  cent. 
This  is  undoubtedly  accounted  for  by  the  fact  that  the  differences  be- 
tween competing  strains  were  so  much  less  than  in  the  oats  variety 
test.  When  the  three  strains  taxonomically  unlike  Red  Rustproof  are 
eliminated,  leaving  12  strains  of  the  same  variety,  the  average  devia- 
tion of  border  yields  from  interior  yields  is  reduced  to  4.69  per  cent 
and  the  average  coefficient  of  competition  to  8.69  per  cent.  It  is  note- 
worthy that  the  competition  between  these  strains  of  the  same  va- 
riety is  decidedly  less  than  that  between  different  varieties.  No  sig- 


Tabee  12. — Correlation  oe  Competition  With  Various  Characteristics  in 

Oats  Strain  Test  1919. 


Character 


Date  of  heading 
Date  of  maturity 
Grain-straw  ratio 
Yield 


Mean  difference  be-  Coefficient  of  correlation 

tween  competing  with  competition 

strains 


2.67  days 
1.56  days 
14.2% 
17.1% 


—.376  ±.136 
— .244  ±.149 
+ .012  ±.159 
+.316  ±.143 


nificant  correlation  was  found  between  these  minor  effects  of  compe- 
tition (for  the  15  strains)  and  the  relative  time  of  heading,  time  of 
maturity,  grain-straw  ratio  or  yield,  as  is  shown  in  Table  12,  though 
in  this  case  again  the  early  strains  and  the  high-yielding  strains  showed 
some  tendency  to  profit  by  competition. 


Experiments  in  Field  Plot  Technic 


37 


In  the  wheat  variety  test  of  1920  the  average  yield  of  border  rows 
differed  from  the  average  yield  of  interior  rows  by  12.30  per  cent  and 
the  mean  coefficient  of  competition  was  19.79  per  cent.  These  figures 
represent  the  average  determinations  when  the  two  varieties  of  rye 
and  the  border  yields  of  the  varieties  of  wheat  adjoining  them  were 
eliminated.  The  correlation  between  competition  and  relative  yield, 
date  of  heading,  and  date  of  maturity  were  determined  for  this  test 
and  the  coefficients  of  correlation  are  shown  in  Table  13. 


Table  13. — Correlation  of  Competition  With  Various  Characteristics  in 
Wheat  Variety  Test  1920. 


Character 


Mean  difference  be-  Coefficient  of  correlation 

tween  competing  with  competition 

varieties 


Date  of  heading  2.3  days  — .515  ±.048 

Date  of  maturity  2.7  days  — .552  ±.045 

Yield  28.9%  +.582  ±.043 


Competition  in  this  test  was  negatively  correlated  with  earliness  of 
heading  and  maturity  and  positively  with  yield.  All  of  the  correla- 
tion coefficients  are  clearly  significant.  In  other  words,  there  was  a 
rather  pronounced  tendency  for  the  early  and  high-yielding  varieties 
to  profit  in  competition.  To  a considerable  extent  the  early  varieties 
were  the  high  yielding  varieties  in  this  test,  as  indicated  by  the  fact 
that  the  correlation  coefficient  for  date  of  heading  and  yield  was  —.511 
±.051,  and  that  for  date  of  maturity  and  yield  was  —.642  ±.041.  Al- 
though it  is  clear  from  these  results  that  early,  high-yielding  varieties 
excelled  in  competition,  it  is  not  clear  whether  they  did  so  chiefly  as  a 
result  of  their  earliness  or  chiefly  as  a result  of  their  yield. 


Table  14. — Correlation  of  Competition  With  Various  Characteristics  in 
Wheat  Variety  Test  1921. 

Character  Mean  difference  be-  Coefficient  of  correlation 

tween  competing  with  competition 

varieties 


Date  of  heading 
Date  of  maturity 
Height 
Yield 


2.1  days 
1.6  days 
3.3  inches 
19.5% 


—.271  ±.060 
—.222  ±.062 
+ .347  ±.057 
+ .294  ±.059 


Similar  results  were  obtained  in  the  wheat  variety  test  of  1921 
in  which  the  difference  between  the  average  yield  of  border  and  in- 
terior rows  was  12.89  per  cent  and  the  mean  coefficient  of  competition 


38 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


was  18.85  per  cent.  Correlations  were  determined  for  competition  and 
relative  yield,  date  of  maturity,  date  of  heading,  and  height  in  this 
test.  The  coefficients  of  correlation  thus  determined  are  shown  in 
Table  14. 

In  this  case,  as  in  the  wheat  variety  test  of  the  preceding  season, 
dates  of  heading  and  maturity  were  correlated  negatively  and  yield 
was  correlated  positively  with  competition.  The  coefficients  of  correla- 
tion were  materially  lower,  and  in  fact  are  hardly  significant.  It  is  in- 
teresting that  in  this  case  height  was  correlated  more  closely  with 
competition  than  were  either  earliness  or  yield.  In  this  season  again 
earliness  was  correlated  to  some  extent  with  yield,  the  coefficients  of 
correlation,  for  date  of  heading  and  yield  being  — .331  ±.062  and  for 
date  of  maturity  and  yield  —.419  ±.057. 

In  the  wheat  mixture  test  of  1921  the  varieties  were  grouped 
roughly  in  respect  to  earliness,  and  in  only  three  cases  was  there  a 
greater  difference  than  two  days  in  heading  or  maturity  between  ad- 
jacent varieties.  The  rows  in  this  test  ran  north  and  south.  The 
conditions  may  be  considered  favorable  in  this  test  for  the  reduction 
of  competition.  Nevertheless  the  average  yield  of  border  rows  dif- 
fered from  that  of  interior  rows  by  10.07  per  cent  and  the  mean  co- 
efficient of  competition  was  14.28  per  cent.  The  coefficients  of  correla- 
tion determined  for  competition  and  date  of  heading,  date  of  maturity, 
and  yield,  are  shown  in  Table  15. 

Table  15. — Correlation  of  Competition  With  Various  Characteristics  in 
Wheat  Mixture  Test  1921. 

Character  Mean  difference  be-  Coefficient  of  correlation 

tween  competing  with  competition 

varieties 

Date  of  heading  1.2  days  — .514  ±.083 

Date  of  maturity  0.8  days  — .613  ±.070 

Yield  19.2%  +.554  ±.078 

In  this  test  significant  negative  correlations  between  competition 
and  dates  of  heading  and  maturity  and  a significant  positive  correla- 
tion between  competition  and  yield  are  shown.  The  tendency  for 
early,  high-yielding  varieties  to  profit  by  competition  was  about  as 
strong  as  in  the  wheat  variety  test  of  the  preceding  season,  though  the 
extent  of  competitive  effect  was  considerably  reduced. 

The  effects  of  competition  in  the  oats  variety  test  in  1921  were 
extreme.  The  yields  of  border  rows  differed  by  16.74  per  cent,  on 
the  average,  from  the  yields  of  interior  rows,  and  the  mean  coefficient 
of  competition  was  39.15  per  cent.  The  extreme  effects  of  compe- 


Experiments  in  Field  Plot  Technic 


39 


tition  in  this  test  are  probably  accounted  for  by  the  fact  that  the  va- 
rieties differed  very  widely  in  varietal  type  and  in  yield.  Differences 
of  as  much  as  17  days  in  date  of  heading,  13  days  in  date  of  maturity, 
and  almost  200  per  cent  in  yield,  were  involved.  The  correlations  de- 
termined between  competition  and  relative  date  of  heading,  date  of 
maturity  and  yield,  are  shown  in  Table  16. 

Table  16. — Correlation  of  Competition  With  Various  Characteristics  in 

Oats  Variety  Test  1921. 

Character  Mean  difference  be-  Coefficient  of  correlation 

tween  competing  with  competition 

varieties 

Date  of  heading  4.8  days  — .648  ±.060 

Date  of  maturity  4.1  days  — .860  ±.028 

Yield  51.1%  +.484  ±.082 

A remarkably  high  negative  correlation  between  date  of  maturity 
and  competition  is  shown.  The  negative  correlation  between  date  of 
heading  and  yield  is  also  quite  high,  while  the  positive  correlation  be- 
tween yield  and  competition  is  barely  significant.  In  this  test,  in  which 
extreme  differences  in  time  of  maturity  occurred,  the  early-maturing 
varieties  had  a very  distinct  advantage  in  competition  with  the  later 
varieties.  Earliness  was  very  closely  correlated  with  yield  in  the  oats 
variety  test  of  this  season,  the  coefficient  of  correlation  for  date  of 
heading  and  yield  being  —.750  =+052  and  that  for  date  of  maturity  and 
yield  being  —.894  ±.024.  Considering  the  close  correlation  of  earliness 
and  yield,  and  the  relatively  low  correlation  of  yield  and  competition, 
it  would  seem  that  the  latter  may  be  merely  a by-product  of  the  rela- 
tion of  earliness  to  competition.  Since  the  early  varieties  were  the 
leaders  both  in  competition  and  in  yield,  some  correlation  of  yield  and 
competition  is  inevitable. 

In  the  oats  strains  test  of  1921  Kherson  and  Red  Rustproof  strains 
were  alternated  and  both  a Kherson  and  a Red  Rustproof  check  were 
grown.  In  most  cases  therefore  the  competing  border  rows  repre- 
sented these  two  varieties,  though  in  some  cases  two  Red  Rustproof 
or  two  Kherson  plots  occurred  together,  as  is  shown  in  the  planting 
plan  in  figure  5.  The  effects  of  competition  in  this  plot  were  quite 
distinct,  as  is  to  be  expected,  though  they  were  not  so  extreme  as  in 
the  oats  variety  test  discussed  above,  which  was  located  on  the  same 
field.  The  average  yield  of  border  rows  differed  from  the  average  yield 
of  interior  rows  by  11.76  per  cent.  The  mean  coefficient  of  compe- 
tition was  23.85  per  cent. 

When  we  exclude  the  competition  between  the  three  strains  not 
true  to  name  and  the  strains  adjacent  to  each,  that  between  the  Kher- 


40 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


son  and  Red  Rustproof  check  plots,  and  that  between  adjacent  strains 
of  the  same  variety,  58  cases  of  competition  between  different  strains 
of  Kherson  and  Red  Rustproof  remain.  In  these  the  mean  yield  of 
border  rows  differed  from  that  of  interior  rows  by  14.06  per  cent  and 
the  mean  coefficient  of  competition  was  30.86  per  cent.  In  every  case 
the  Kherson  strain  outyielded  the  adjacent  Red  Rustproof  strain, 
though  the  advantage  in  yield  varied  from  27  per  cent  to  165  per  cent. 
Similarly,  the  Kherson  strains  were  earlier  in  maturity  and  heading, 
and  taller,  in  each  case,  but  with  a rather  wide  variation  in  the  extent 
of  their  advantage.  In  all  but  three  of  the  58  cases  the  Kherson  strains 
showed  a greater  advantage  in  yield  over  the  adjacent  Red  Rustproof 
strains  in  their  competing  border  rows  than  in  their  interior  rows. 
The  average  yields  of  the  30  Red  Rustproof  strains  and  29  Kherson 
strains,  in  interior  rows  and  competing  border  rows,  were  as  follows : 


Average  yield  in 

Average  yield  in 

interior 

rows 

competing  border 

rows 

Bushels 

Relative 

Bushels 

Relative 

Red  Rustproof  strains 

21.00 

100 

18.59 

100 

Kherson  strains 

j 35.79 

170 

41.00 

222 

The  Kherson  strains  outyielded  the  Red  Rustproof  strains  by  70 
per  cent  in  their  interior  rows  and  by  122  per  cent  in  their  competing 
border  rows.  The  coefficients  of  competition,  like  the  relative  yield, 
earliness,  and  height,  varied  rather  widely.  Correlations  were  there- 
fore measured  for  the  advantage  of  the  Kherson  strain  of  each  adjacent 
pair  in  competition  and  its  advantages  in  yield,  date  of  heading,  date 
of  maturity,  and  height.  The  coefficient  of  correlation  in  each  case 
was  insignificant. 

Discussion. — In  each  of  these  tests,  with  the  exception  of  the 
oats  strain  test  of  1919,  in  which  most  of  the  strains  compared  be- 
longed to  the  same  variety,  border  rows  differed  from  interior  rows  in 
yield  by  more  than  10  per  cent.  Differences  as  great  as  this  will  change 
materially  the  relative  standing  of  varieties.  In  single-row  tests  the 
effects  of  competition  would  be  considerably  greater  than  in  these 
border  rows,  affected  by  competition  on  only  one  side.  Furthermore, 
in  each  test,  of  course,  there  were  many  cases  in  which  competition 
caused  much  larger  differences  in  yield  than  are  shown  by  average 
figures. 


Experiments  in  Field  Plot  Technic 


41 


The  relation  of  the  direction  of  rows  to  the  effects  of  varietal  com- 
petition is  not  clearly  shown  by  these  experiments.  The  tests  which 
showed  least  the  effect  of  competition,  the  oats  strain  test  of  1919 
and  the  wheat  mixture  test  of  1921,  were  in  rows  running  north  and 
south.  But  relatively  little  effect  from  competition  is  to  be  expected 
in  these  tests,  regardless  of  the  direction  of  the  rows,  because  of  the 
similarity  of  adjacent  strains.  In  the  oats  strain  test  12  of  the  15 
strains  were  taxonomically  identical,  and  it  has  been  shown  that  the 
effects  of  competition  among  these  was  much  less  than  among  the 
strains  of  different  varieties.  In  the  wheat  mixture  test  the  varieties 
making  up  each  mixture,  which  were  grown  side  by  side  in  the  test, 
were  chosen  partly  for  their  similarity  in  time  of  maturity,  and  the 
differences  between  adjacent  varieties  were  therefore  considerably 
less  than  in  the  wheat  variety  test  of  the  same  season.  It  cannot  be 
stated  definitely,  therefore,  from  the  results  of  these  tests,  that  tests 
in  rows  running  north  and  south  are  either  more  or  less  subject  to  error 
from  competition  than  tests  in  rows  running  east  and  west.  It  is 
clear,  however,  that  a considerable  error  from  varietal  competition 
may  occur  in  tests  in  which  the  rows  run  north  and  south,  as  is  evi- 
denced particularly  by  the  barley  and  oats  variety  tests  of  1919. 

The  relation  of  competition  to  relative  date  of  heading,  date  of 
maturity,  grain-straw  ratio,  height,  and  yield,  insofar  as  it  was  in- 
vestigated in  these  experiments,  is  shown  in  summary  form  in  Table 
17.  Although  none  of  these  characteristics  shows  a significant  rela- 


Tabee  17. — Summary  of  Effects  of  Competition  in  Aee  Tests. 


Test 

Season .... 

Number  of 
varieties  or 
strains j 

Mean  coeffi- 
cient of 
competition. . . 

Date  of 
Heading. 

Coefficient  of 

Correlation  between  Competition  and — 

Date  of 
Maturity. 

Grain- Straw 
Ratio. 

Height. 

Yield. 

Barley  variety 

1919 

| 27 

21.30 

— .153  ± . 120 

— .063±.123 

+ .072  + .122 

+ .442±.099 

Oats  variety 

1919 

24 

27.67 

— .456±.103 

— .091  + . 129 

-f-.314-H.H7 

Oats  strain 

1919 

15 

13.11 

-.376±.136 

— .244±.157 

+ .012  + .159 

+ .316+.143 

Wheat  variety 

1920 

1 94 

19.79 

— .5l5±.048 

— .552  + . 045 

+ .582  + . 043 

Wheat  variety 

1921 

94 

18.85 

— .271±.060 

— .222  + . 062 

+ .347±.057 

+ .294  + .059 

Wheat  mixture 

1921 

1 30 

14.28 

— .514  + . 083 

— .613±.070 

+ .554±.078 

Oats  variety 

1921 

32 

39.15 

— .648  + .060 

— .860±.028 

+.484  + .082 

tion  to  competition  in  every  case,  the  results  of  the  tests  are  fairly 
consistent.  The  correlation  of  competition  with  yield  is  always  posi- 
tive, and  is  fairly  high  in  every  case,  the  lowest  coefficient  being  -(—.294 
—.059.  From  these  results  there  can  be  no  doubt  that  the  higher  yield- 
ing varieties  are  those  which  in  general  have  profited  by  competition. 
The  date  of  heading  and  the  date  of  maturity  show  a negative  correla- 


42 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


tion  with  competition  in  each  case,  though  some  of  the  coefficients 
are  insignificant.  It  is  clear  therefore  that  early  varieties  are,  in  gen- 
eral, able  to  compete  more  strongly,  but  the  extent  of  this  relation  is 
quite  variable.  The  grain-straw  ratio  showed  no  significant  relation 
to  competition  in  any  of  the  experiments  of  1919,  and  was  not  deter- 
mined for  the  succeeding  tests.  Height  was  correlated  positively  with 
competition  in  the  one  test  in  which  height  was  determined,  the  wheat 
variety  test  of  1921.  In  this  test  height  was  more  closely  related  to 
competition  than  were  date  of  heading,  date  of  maturity,  or  yield. 

In  the  oats  variety  tests,  the  relation  of  early  maturity  to  compe- 
tion  is  particularly  marked,  the  coefficients  of  correlation  in  both  oats 
variety  tests  being  distinctly  greater  for  date  of  maturity  and  compe- 
tition than  for  yield  and  competition.  In  the  wheat  tests  there  was 
little  difference  in  the  degree  of  relation  to  competition  between  earli- 
ness and  yield.  In  the  one  test  of  barley  varieties  conducted,  yield 
was  more  closely  correlated  with  competition  than  was  either  the 
date  of  heading  or  date  of  maturity,  but  none  of  the  three  showed  a 
clearly  significant  correlation. 

It  is  clear  that  in  these  trials  the  early,  high-yielding  varieties 
profited  by  competition.  To  a considerable  extent  these  may  be  the 
same  varieties,  for  the  correlation  of  earliness  and  yield  was  high  in 
most  of  the  tests  conducted.  The  relation  of  earliness  and  other 
characters  to  yield  under  Missouri  conditions  will  be  considered  more 
fully  in  another  paper,  but  data  of  interest  in  this  connection  are  ap- 
propriate here.  The  coefficients  of  correlation  of  yield  with  date  of 
heading  and  date  of  maturity  in  the  variety  tests  discussed  in  this 
paper  are  shown  in  Table  18. 

Table  18. — Correlation  of  Yield  With  Dates  of  Heading  and  Maturity  in 
Variety  Tests  of  Barley,  Oats,  and  Wheat 

Coefficient  of  correlation  of 
Number  of  yield  with — 


Crop 

Season 

varieties 

Date  of  heading 

Date  of  maturity 

Barley 

1919 

27 

—.281  ±.120 

—.271  ±.120 

Oats 

1919 

40 

—.627  ±.065 

Oats 

1921 

32 

— .750  ±.052 

— .894  ±.024 

Wheat 

1920 

94 

— .511  ±.051 

— .642  ±.041 

Wheat 

1921 

94 

—.331  ±.062 

—.419  ±.057 

When  a very  high  correlation  exists  between  earliness  and  yield 
it  is  likely  that  a character  closely  correlated  with  one  may  show  a 
high  degree  of  correlation  with  the  other,  which  might  not  be  shown 
were  it  not  for  the  first  correlation.  For  example,  suppose  earliness 


Experiments  in  Field  Plot  Technic 


43 


of  maturity  is  largely  responsible  for  strong  competitive  value.  Then 
in  a season  when  earliness  is  closely  correlated  with  yield  a close  cor- 
relation of  competition  and  yield  is  likely  to  be  found,  not  because  high 
yield  makes  for  strong  competition  but  because  the  high-yielding  va- 
rieties are  early.  Conversely,  the  competing  value  may  be  dependent 
on  the  yield  and  the  correlation  with  earliness  may  be  incidental,  under 
the  same  conditions.  If  the  relation  of  earliness  and  yield  were  con- 
stant, such  a question  would  have  little  practical  importance,  but  when 
the  relation  is  reversed,  as  it  may  be  in  different  localities  and  even  in 
different  seasons  in  the  same  locality,  the  relation  of  competition  to 
the  two  characteristics  may  be  very  different.  The  relation  of  compe- 
tition to  earliness  and  yield  in  these  tests,  therefore,  may  be  due  pri- 
marily to  the  predominating  influence  of  either  of  these  two  charac- 
teristics, or  to  the  influence  of  both. 

General  conclusions  regarding  competition  should  not  be  drawn 
from  these  tests.  The  problem  of  competition  is  complicated  by  many 
factors,  and  will  require  numerous  and  extensive  investigations  for 
its  solution.  These  results,  however,  indicate  that  gross  errors  from 
this  source  are  commonly  involved  in  variety  tests,  that  such  errors 
occur  both  in  rows  running  east  and  west  and  in  rows  running  north 
and  south,  that  the  error  is  less  when  the  varieties  and  strains  com- 
pared are  structurally  similar  than  when  they  are  widely  different,  and 
that  the  error  may  be  reducible  to  some  extent  by  the  grouping  of  va- 
rieties according  to  the  time  of  maturity  and  possibly  other  characters, 
when  the  relation  of  such  characters  to  competition  is  more  fully 
studied.  In  the  present  state  of  knowledge  regarding  the  relation  of 
competition  to  the  characteristics  of  the  varieties  compared,  the  use 
of  border  rows  is  highly  desirable,  since  by  their  use  the  error  from 
competition  can  be  practically  eliminated. 

SIZE  AND  REPLICATION  OF  PLOTS. 

Previous  Investigation. — Most  of  the  direct  evidence  reported 
on  replication  and  size  of  plots  has  been  obtained  in  experiments  in 
which  a field  of  a uniformly  handled  crop  is  harvested  in  a large  num- 
ber of  small  sections.  These  sections  are  grouped  to  form  plots  of 
different  shapes  and  sizes,  and  systematically  distributed  sections  are 
averaged  to  represent  replicate  plots.  The  relative  variability  of  the 
yields  determined  by  each  plot  arrangement  is  the  criterion  of  expe- 
rimental accuracy.  Such  experiments  have  been  reported  by  Morgan15 
with  wheat  and  fodder  corn,  Wood  and  Stratton18  with  mangels,  Mer- 
cer and  Hall  12  with  wheat  and  mangels,  Hall  and  Russell3  with  wheat, 


44 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


Montgomery13’14  with  wheat,  Kiesselbach5  with  oats,  and  Day1  with 
wheat. 

The  general  conclusions  drawn  from  these  experiments  are  in 
harmony,  though  the  specific  size  and  shape  of  plot  and  number  of 
replications  found  most  desirable  vary  rather  widely.  In  general,  plot 
variability  was  reduced  by  increasing  the  size  of  the  individual  plot, 
up  to  a certain  limit,  but  it  was  reduced  much  more  effectively  by  rep- 
lication of  plots.  For  a given  area  a large  number  of  small  plots  was 
always  found  more  accurate  than  a small  number  of  large  plots. 

But  the  size  of  the  plot  cannot  be  reduced  indefinitely  for  several 
reasons.  As  the  plot  becomes  smaller  the  proportion  subject  to  “bor- 
der effect”  rapidly  becomes  greater.  This  border  effect  may  be  due 
to  the  modified  growth  of  plants  adjoining  an  alley  or  to  the  in- 
fluence of  the  competition  of  different  varieties  in  adjacent  rows.  If 
the  borders  are  not  discarded  an  important  systematic  error  is  involved ; 
if  they  are  discarded  a considerable  portion  of  the  land  and  labor  is 
lost.  In  either  case  the  disadvantage  is  increased  as  the  size  of  the 
plot  is  decreased.  When  single  rod-row  plots  ar,e  used  the  whole 
plot  is  subject  to  border  effect.  The  importance  of  this  error  has 
already  been  discussed.  Another  disadvantage  of  the  extremely 
small  plot  is  that  slight  differences  in  stand  and  small  mechanical  er- 
rors have  a marked  effect  on  the  yields.  The  increased  labor  involved 
in  handling  a large  number  of  small  plots  rather  than  a small  number 
of  large  plots  is  also  an  important  disadvantage. 

The  length  of  the  so-called  rod-row  has  usually  been  determined 
by  convenience.  Commonly  used  lengths  when  the  rows  are  a foot 
apart  are  16  feet  for  wheat,  20  feet  for  barley,  and  15  feet  for  oats, 
since  with  these  lengths  yields  in  grams  per  row  may  easily  be  con- 
verted to  bushels  per  acre.  In  other  cases  the  most  convenient  length 
is  determined  by  the  dimensions  of  experiment  fields.  Although  in- 
creasing the  length  of  the  row  would  doubtless  reduce  variability,  a 
greater  gain  could  be  made  on  the  same  area  by  further  replication. 
Ordinarily  it  is  preferable,  therefore,  to  retain  the  most  convenient 
length  and  to  make  any  desired  increase  in  size  of  plot  in  the  width, 
for  widening  the  plots  will  rapidly  reduce  the  proportion  subject  to 
border  effect. 

Experimental  Results.  —Size  of  Plots. — By  comparing  the  stand- 
ard deviations  of  single  rows  and  blocks  consisting  of  three  and  five 
rows  each,  in  the  check  plots,  it  is  possible  to  determine  the  relative 
value  of  plots  of  the  three  sizes  in  counteracting  plot  variability.  In 
this  comparison  the  single-row  and  three-row  plots  correspond  respec- 
tively to  3-row  and  5-row  plots  in  which  the  border  rows  are  dis- 


Experiments  in  Field  Plot  Technic 


45 


carded,  since  they  are  made  up  of  rows  protected  from  varietal  compe- 
tition by  border  rows.  In  each  of  the  computations  summarized  be- 
low each  check  plot  is  represented  by  only  one  yield.  For  example,  in 
determining  the  yield  and  standard  deviation  of  single  rows  in  the  20 
check  plots  of  the  oats  variety  test  of  1919,  the  constants  for  single 
rows  are  the  average  of  determinations  made  independently  for  Row 
2 of  each  of  the  20  plots,  for  Row  3,  and  for  Row  4.  The  determina- 
tions for  3-row  plots  are  similarly  made  from  the  computed  yields  of 
the  three  interior  rows  of  each  check  plot,  and  those  for  5-row  plots 
from  the  computed  yields  of  the  entire  plots.  Thus  each  determination 
represents  the  same  number  of  plots  and  the  same  area,  the  only  dif- 
ference being  in  the  size  of  the  individual  plot.  It  would  be  possible, 
of  course,  to  test  40  per  cent  more  varieties  with  the  same  number  of 
replications  or  to  increase  the  number  of  replications  by  40  per  cent 
for  the  same  number  of  varieties  on  the  same  area,  if  3-row  blocks 
were  used  rather  than  5-row  blocks. 

The  yield  and  variability  of  check  plots  of  different  sizes  in  the 
barley  variety  test  of  1919  are  shown  in  Table  19.  The  variety  grown 
in  these  check  plots  was  Oderbrucker,  seeded  at  the  rate  of  8 pecks 
per  acre.  The  check  variety  was  grown  in  every  sixth  plot. 


Table  19. — Yield  and  Variability  of  Check  Plots. 
Single-row,  Three-row,  and  Five-row — Barley  Variety  Test  1919. 


Size  of  plot 

Number 
of  plots 

Yield 
per  acre 

Standard  deviation 

bu. 

bu. 

% 

Single-row 

Row  1 

21 

41.26 

7.95 

19.26 

Row  2 

21 

36.71 

8.30 

22.61 

Row  3 

21 

37.15 

8.48 

22.84 

Row  4 

21 

35.82 

10.37 

28.96 

Row  5 

21 

42.12 

11.86 

28.16 

Mean  of  three 

interior  rows 

21 

36.56 

9.05 

24.80 

Mean  of 

five  rows 

21 

38.61 

9.39 

24.37 

Three-row  Plot 

(Interior  rows) 

21 

36.56 

8.11 

22.18 

Five-row  Plot 

21 

38.61 

8.29 

21.47 

The  variability  of  the  single-row  plots  is  12  per  cent  higher  on 
the  average  than  that  of  the  3-row  plots.  That  is,  3-row  plots  with 


46 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


borders  discarded  would  have  given  in  this  case  somewhat  more  va- 
riable results  than  5-row  blocks  with  borders  discarded.  The  same  5- 
row  blocks  harvested  entire  (with  borders  retained)  gave  slightly  less 
variable  yields  than  when  the  borders  were  discarded. 

The  same  comparison  may  be  made  in  the  check  plots  of  the 
oats  variety  test  of  1919.  The  check  variety  was  Red  Rustproof, 
drilled  at  the  rate  of  10  pecks  per  acre  in  every  ninth  plot.  The  re- 
sults are  shown  in  Table  20. 


Table  20. — Yield  and  Variability  of  Check  Plots. 
Single-row,  Three-row,  and  Five-row — Oats  Variety  Test  1919. 


Size  of  plot 

Number 
of  plots 

Yield 
per  acre 

Standard  deviation 

bu. 

bu. 

% 

Single-row 

Row  1 

20 

44.64 

10.37 

23.23 

Row  2 

20 

47.97 

9.81 

20.45 

Row  3 

20 

46.56 

11.42 

24.53 

Row  4 

20 

46.95 

13.81 

29.41 

Row  5 

20 

42.09 

11.58 

27.51 

Mean  of  three 

interior  rows 

20 

47.16 

11.68 

24.80 

Mean  of 

five  rows 

20 

45.64 

11.40 

25.03 

Three-row  Plot 

(Interior  rows) 

20 

47.16 

10.62 

22.59 

Five-row  Plot 

20 

45.64 

9.72 

21.30 

The  results  in  this  case  are  practically  identical  with  those  of  the 
barley  variety  test.  Protected  single  rows  were  10  per  cent  more  va- 
riable than  protected  3-row  blocks,  while  the  latter  were  only  6 per 
cent  more  variable  than  unprotected  5-row  blocks. 

In  the  test  of  strains  of  Red  Rustproof  oats,  conducted  on  the 
same  field  in  1919,  adjoining  the  oats  variety  test,  the  same  variety 
was  used  as  check,  and  the  crop  was  seeded  on  the  same  day  with  the 
same  machine,  but  the  check  plots  were  in  every  sixth  instead  of  every 
ninth  plot.  The  corresponding  data  for  these  check  plots  are  given  in 
Table  21. 

Although  the  variability  of  these  plots  is  lower,  the  relative  va- 
riability of  plots  of  different  sizes  is  similar  to  that  of  the  variety  test. 
The  single  interior  rows  are  on  the  average  24  per  cent  more  variable 
than  the  3-row  block.  The  3-row  plot  is  only  very  slightly  more  va- 
riable than  the  5-row  plot. 


Experiments  in  Fieed  Plot  Technic 


47 


Table  21. — Yield  and  Variability  of  Check  Plots. 
Single-row,  Three-row,  and  Five-row. — Oats  Strain  Test  1919. 


Size  of  plot 

Number 
of  plots 

Yield 
per  acre 

Standard  deviation 

bu. 

bu. 

% 

Single-row 

Row  1 

18 

41.87 

6.35 

15.15 

Row  2 

18 

40.88 

5.52 

13.51 

Row  3 

18 

43.50 

5.81 

13.37 

Row  4 

18 

45.00 

7.31 

16.25 

Row  5 

18 

41.50 

6.37 

15.35 

Mean  of  three 

interior  rows 

18 

43.13 

6.21 

14.38 

Mean  of 

five  rows 

18 

42.55 

6.27 

14.73 

Three-row  Plot 

(Interior  rows) 

18 

43.13 

5.04 

11.68 

Five-row  Plot 

18 

42.55 

4.86 

11.41 

In  the  wheat  variety  test  of  1920  the  check  variety  was  Fultz, 
which  was  seeded  at  the  rate  of  six  pecks  per  acre  in  every 
seventh  plot.  The  results  of  interest  in  this  connection  are  shown  in 
Table  22. 


Table  22. — Yield  and  Variability  of  Checks  Plots. 
Single-row,  Three-row,  and  Five-row. — Wheat  Variety  Test  1920 


Size  of  plot 

Number 
of  plots 

Yield 
per  acre 

Standard  deviation 

bu. 

bu. 

% 

Single-row 

Row  1 

80 

20.74 

6.58 

31.72 

Row  2 

80 

17.28 

5.02 

29.05 

Row  3 

80 

18.34 

4.50 

24.52 

Row  4 

80 

17.29 

5.10 

29.48 

Row  5 

80 

19.37 

6.00 

30.97 

Mean  of  three 

interior  rows 

80 

17.64 

4.87 

27.68 

Mean  of 

five  rows 

80 

18.60 

5.44 

29.15 

Three-row  Plot 

(Interior  rows) 

80 

17.64 

4.43 

25.11 

Five-row  Plot 

80 

18.63 

4.77 

25.60 

48  Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 

Again  the  single  rows  are  distinctly  more  variable  than  the  3-row 
plot,  in  this  case  to  the  extent  of  10  per  cent.  The  5-row  and  the 
3-row  plots  are  about  equally  variable,  the  slight  advantage  in  this  case 
being  in  favor  of  the  latter. 

To  summarize,  it  is  evident  that  the  protected  3-row  plot  is  some- 
what less  subject  to  plot  variability  than  the  protected  single-row,  but 
the  relative  value  of  the  5-row  plot  harvested  entire  and  the  same  plot 
harvested  as  a protected  3-row  block  is  not  clear.  Some  further 
comparison  of  these  two  methods  was  made  in  1921.  The  variability 
of  the  check  plots  in  both  the  wheat  and  oats  tests  was  computed  as 
protected  3-row  and  as  unprotected  5-row  plots.  In  the  wheat  tests 
the  check  variety  was  Poole,  seeded  at  5 pecks  per  acre  in  every 
seventh  plot  in  the  variety  test,  and  in  every  sixth  plot  in  the  mixture 
test.  In  the  oats  tests  the  check  variety  was  Kherson,  seeded  at  10 
pecks  per  acre  in  every  sixth  plot.  The  results  are  shown  in  Table  23. 


Table  23. — Yield  and  Variability  of  Check  Plots. 
Three-row  and  Five-row. — Wheat  and  Oats  Tests,  1921. 


Size  of  plot 

Number 
of  plots 

Yield  j 
per  acre  j 

Standard  deviation 

Wheat  Variety  Test 

Three-row  Plots 

80 

bu. 

14.89 

bu. 

2.16 

% 

14.50 

(Interior  rows) 
Five-row  Plots 

80 

13.98 

1.90 

13.61 

Wheat  Mixture  Test 

Three-row  Plots 

30 

15.48 

3.25 

20.98 

(Interior  rows) 
Five-row  Plots 

30 

15.78 

3.55 

22.49 

Oats  Variety  and  Strain  Tests 

Three-row  Plots 

120 

37.95 

4.61 

12.15 

(Interior  rows) 
Five-row  Plots 

120 

38.37 

4.70 

12.25 

In  no  case  are  the  differences  very  great.  The  variability  of  3-row 
blocks  is  slightly  greater  in  the  mixture  test  and  that  of  5-row  blocks 
in  the  variety  test  of  wheat.  There  is  practically  no  difference  between 
the  two  in  the  oats  tests. 

Apparently  there  is  no  constant  material  gain  in  plot  uniformity 
obtained  by  the  inclusion  of  the  border  rows  of  the  5-row  plot,  even 
though  the  size  of  the  plot  is  materially  increased  by  this  procedure. 
Even  if  variability  were  decreased  by  their  inclusion,  the  practice  would 
be  of  doubtful  value  in  most  tests,  for  the  reasons  given  in  the  last 
section ; but  with  practically  no  decrease  in  variability  there  is  left  no 


Experiments  in  Field  Plot  Technic 


49 


reason  for  the  harvesting  of  these  rows.  They  are  not  wasted  because 
they  are  not  harvested,  for  they  serve  a valuable  purpose;  the  waste 
would  be  involved  rather  in  harvesting  them,  for  the  added  labor  and 
expense  would  contribute  nothing  to  the  accuracy  of  the  experiment. 

Although  protected  3-row  plots  are  less  variable  than  protected 
single-row  plots,  they  are  not  necessarily  preferable.  Three  protected 
3-row  plots  require  the  same  area  as  five  protected  single-row  plots, 
and  the  harvesting  of  almost  twice  as  large  a crop  (nine  rows  in  the 
first  case  for  every  five  in  the  second).  If  the  mean  yield  of  five 
single  rows  has  as  low  a probable  error  as  the  mean  yield  of  three 
3-row  plots,  the  protected  single-row  plot  will  ordinarily  be  pre- 
ferable, because  of  the  reduction  of  labor  in  harvesting  and  thresh- 
ing. When  the  standard  deviation  of  the  check  plot  yields  is  known, 
the  probable  error  of  the  mean  of  any  number  of  replicate  plots  can 
be  computed  and  the  number  of  replications  for  any  given  degree  of 
accuracy  determined.  If  single-row  plots  were  29  per  cent  more 
variable  than  3-row  plots,  the  probable  errors  of  the  mean  of  three 
3-row  plots  and  of  five  single-row  plots  would  be  equal,  since  the  prob- 
able error  of  the  mean  is  equal  to  the  probable  error  of  a single  deter- 
mination divided  by  the  square  root  of  the  number  of  determinations, 
and  since  the  square  root  of  5 is  29  per  cent  greater  than  the  square 
root  of  3.  In  the  cases  herein  cited  the  advantage  of  the  3-row  plots 
was  considerably  less  than  29  per  cent  in  every  case,  and  we  may  con- 
fidently expect  therefore  that  protected  single-row  plots  repeated  five 
times  will  be  less  variable  than  protected  three-row  plots  repeated  three 
times,  which  would  require  the  same  area  and  more  labor. 

Some  further  evidence  on  the  relative  variability  of  the  protected 
3-row  plot  and  the  unprotected  5-row  plot,  or,  in  other  words,  of 
5-row  plots,  harvested  with  and  without  their  border  rows,  may  be  ob- 
tained from  the  yields  of  the  tested  varieties  and  strains.  Since  the 
number  of  replications  of  each  strain  is  small,  average  deviations  are 
given  instead  of  standard  deviations.  The  inclusion  of  border  rows  in 
the  5-row  plots  should  not  increase  variability,  since  the  adjacent  va- 
rieties are  the  same  in  each  series,  and  the  competitive  effect  should  be 
no  more  variable  than  would  be  that  of  the  same  variety.  A clear-cut 
comparison  of  5-row  and  3-row  plots  is  therfore  available  in  this  case. 
In  the  case  of  the  check  plots  this  comparison  was  somewhat  obscured 
by  the  competitive  effect  of  different  varieties  on  the  border  rows,  which 
might  be  expected  to  increase  variability  and  thus  to  conceal  a possible 
advantage  of  the  5-row  plot. 

The  average  variability  of  3-row  and  5-row  plots  in  the  strains 
tested  in  these  experiments  is  shown  in  Table  24.  Tn  each  case  the 


50 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


figure  given  is  the  mean  of  the  average  variabilities  determined  for  all 
of  the  varieties  or  strains  in  the  experiment. 


Table  24. — Yield  and  Variability  oe  Test  Plots. 
Three-row  and  Five-row. 


Test 

Season 

Number 
of  vari- 
eties 

I 

Number  1 
of  Repli-  j 
cations 

- 

Three-row  Plots  1 

Five-row  Plots 

Yield 
bu.  per 
acre 

Average ! 
Devia- 
tion 
% 

Yield 
bu.  per 
acre 

! Aver- 
i age 
; Devia- 
i tion 
% 

Barley  varieties 

1919 

27 

3 

22.06 

15.35 

21.95 

15.44 

Oats  strains 

1919 

15 

4 

50.07 

5.96 

50.20  ! 

5.10 

Wheat  varieties 

1920 

| 96 

4 

13.39 

24.27 

13.78  | 

24.36 

Wheat  varieties 

1921 

96  i 

4 

15.42 

10.30 

15.57 

9.74 

Wheat  mixtures 

1921 

i 30  ! 

4 

17.62 

9.84 

18.15 

10.03 

Oats  varieties 

1921 

32  i 

4 

29.85 

10.86 

30.70  i 

i 10.14 

Oats  strains 

1921 

64  i 

4 

28.40 

10.82 

j 28.63 

10.58 

There  is  no  consistent  difference  in  variability  between  the  3-row 
plots  and  the  5-row  plots.  In  some  cases  the  former  are  more  va- 
riable; in  others  the  latter;  and  in  no  case  is  the  difference  in  varia- 
bility great.  These  results  are  contrary  to  the  general  impression  that 
variability  decreases  with  increase  in  size  of  plots.  Apparently,  in 
tests  of  this  kind,  the  3-row  plot  is  la^ge  enough  to  give  a fair  sample 
and  nothing  is  gained  by  adding  the  other  two  rows.  When  it  is  con- 
sidered that  the  addition  of  these  two  rows  undoubtedly  introduces 
systematic  error  from  competition  to  a greater  or  less  extent,  and 
involves  a very  considerable  increase  in  the  labor  of  harvesting  and 
threshing,  there  remains  little  doubt  that  the  border  rows  of  5-row 
plots  are  best  discarded  in  experiments  of  this  sort. 

Replication  of  Plots . — It  is  generally  considered  that  the  error 
from  soil  variability  may  be  reduced  to  any  desired  point  by  replica- 
tion in  sufficient  degree.  For  any  given  degree  of  precision  the  num- 
ber of  replications  required  is  dependent  on  the  variability  of  the 
replicate  plots.  When  every  plot  in  a single-row  test  is  provided  with 
two  border  rows  the  area  required  for  the  test  is  tripled,  the  replicate 
plots  are  separated  more  widely,  and  variability  is  usually  increased, 
since  the  range  of  soil  variability  will  usually  be  greater  when  a larger 
area  is  included. 

The  removal  of  border  effect  from  the  rows  harvested  for  yield 
may  in  some  cases  reduce  variability  more  than  enough  to  balance  this 
increase,  but  when  the  unprotected  single  rows  are  grown  in  the  same 
order  in  each  series,  variability  will  not  be  much  affected  by  competi- 
tion, as  before  stated.  Consequently  more  replications  of  single-row 


Experiments  in  Field  Plot  Technic 


51 


plots  protected  by  borders  than  of  the  single-row  plots  not  so  pro- 
tected may  actually  be  required  for  a given  degree  of  plot  variability. 
Similarly,  more  replications  may  be  required  in  a test  of  a large  num- 
ber of  strains  than  in  a test  of  a small  number,  as  Montgomery14  has 
suggested. 

The  number  of  replications  required  may  be  determined  with  a 
fair  degree  of  accuracy  from  the  variability  of  the  check  plots.  The 
variability  of  the  check  plots  in  parts  of  the  large  fields  used  as  com- 
pared with  the  variability  of  the  check  plots  in  the  whole  fields  shows 
the  importance  of  this  point.  In  Table  25  are  given  the  standard  de- 


Table  25. — Relation  of  Plot  Variability  to  Size  of  Experiment  Field. 
Check  Plots  in  Wheat  Variety  Test  1920. 


Size  of  field 

No.  of 
Plots 

Yield 

bu.  per  acre 

Standard  deviation 
bu.  % 

Four  ranges  (1st)* 

20 

14.79 

3.789 

25.62 

Four  ranges  (2nd) 

20 

18.35 

4.073 

22.20 

Four  ranges  (3rd) 

20 

16.67 

3.659 

21.94 

Four  ranges  (4th) 

20 

20.74 

3.876 

18.69 

Mean 

20 

17.64 

3.849 

22.11 

Eight  ranges  (1st) 

40 

16.57 

4.316 

26.05 

Eight  ranges  (2nd) 

40 

18.71 

4.285 

22.90 

Mean 

40 

17.64 

4.302 

24.48 

Sixteen  ranges 

80 

17.64 

4.430 

25.11 

♦The  four-range  and  eight-range  sections  are  in  order  from  west  to  east. 


viations  of  the  yields  of  the  check  plots  in  the  wheat  variety  test  of 
1920.  The  yields  of  the  three  interior  rows  of  the  check  plots  were 
used  in  computing  these  constants. 

Twenty-four  varieties  could  have  been  replicated  four  times  in 
the  four  ranges  comprising  any  quarter  of  the  field.  As  the  probable 
error  of  a single  plot  yield  is  14.92  per  cent  we  may  conclude  that  the 
probable  error  of  the  mean  of  four  such  yields  would  be  about  7.46 
per  cent.  But  when  96  varieties  must  be  tested,  as  they  were  in  this 
test,  four  replications  require  16  ranges,  and  the  probable  error  of  the 
mean  yield  becomes  8.47  per  cent.  A degree  of  precision  which  could 
be  attained  with  four  replications  in  a test  covering  four  ranges  could 
hardly  be  attained  with  five  replications  in  a test  covering  sixteen 
ranges. 

Corresponding  data  for  the  wheat  variety  test  of  1921  are  given 
in  Table  26.  Although  the  variability  in  this  experiment  was  much 
lower,  the  relative  variability  of  large  and  small  experiment  fields  was 


52 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


Table  26. — Relation  oe  Plot  Variability  to  Size  oe  Experiment  Field. 
Check  Plots  in  Wheat  Variety  Test  1921. 


Size  of  field 

No.  of 
Plots 

Yield 

bu.  per  acre 

Standard  Deviation 
bu.  % 

Four  ranges  (1st)* 

16 

15.78 

1.584 

10.04 

Four  ranges  (2nd) 

16 

15.48 

1.586 

10.25 

Four  ranges  (3rd) 

16 

15.28 

2.099 

13.74 

Rour  ranges  (4th) 

16 

13.01 

2.091 

16.07 

Mean 

16 

14.89 

1.840 

12.53 

Eight  ranges  (1st) 

32 

15.63 

1.592 

10.19 

Eight  ranges  (2nd) 

32 

14.14 

2.383 

16.85 

Mean 

32 

14.89 

1.988 

13.52 

Sixteen  ranges 

64 

14.89 

2.159 

14.50 

*The  four-range  and  eight-range  sections  are  in  order  from  west  to  east. 


similar.  Again  the  degree  of  accuracy  obtained  with  four  replications 
in  four  ranges  would  have  been  unattainable  with  five  replications  in 
16  ranges. 

The  oats  variety  test  and  strain  test  in  1921  were  contiguous,  oc- 
cupying 24  ranges,  with  120  check  plots  of  Kherson  oats,  or  one  in 
every  sixth  plot.  The  variability  of  these  check  plots  in  sections  of 

Table  27. — Relation  oe  Plot  Variability  to  Size  oe  Experiment  Field. 

Check  Plots  in  Oats  Variety  and  Strain  Test,  1921. 


No.  of 

Size  of  field  plots  Yield  Standard  deviation 

bu.  per  acre  bu.  % 


Four  ranges  (1st) 
Four  ranges  (2nd) 
Four  ranges  (3rd) 
Four  ranges  (4th) 
Four  ranges  (5th) 
Four  ranges  (6th) 
Mean 

Eight  ranges  (1st) 
Eight  ranges  (2nd) 
Eight  ranges  (3rd) 

Mean 

Twelve  ranges  (1st) 
Twelve  ranges  (2nd) 

Mean 

Twenty- four  ranges 


20 

35.81 

20 

34.95 

20 

38.14 

20 

39.60 

20 

38.91 

20 

40.31 

20 

37.95 

40 

35.38 

40 

38.87 

40 

39.60 

40 

37.95 

60 

36.30 

60 

39.60 

60 

37.95 

120 

37.95 

4.75 

13.26 

2.90 

8.30 

4.21 

11.04 

4.66 

11.77 

4.14 

10.65 

4.14 

10.27 

4.13 

10.88 

3.96 

11.19 

4.50 

11.58 

4.21 

10.62 

4.22 

11.13 

4.25 

11.71 

4.36 

11.01 

4.31 

11.36 

4.61 

12.15 

Experiments  in  Field  Plot  Technic 


53 


four,  eight,  and  twelve  ranges,  and  in  the  whole  field  of  24  ranges,  is 
shown  in  Table  27. 

The  variability  of  the  whole  field  of  24  ranges  was  12  per  cent 
greater  than  the  average  variability  of  sections  of  four  ranges  each. 
In  this  case  again,  five  replications  in  the  larger  field  would  have  given 
less  accurate  results  than  four  replications  in  the  smaller. 

In  each  of  the  cases  cited  above  a steady  increase  in  variability 
is  apparent  as  the  size  of  the  experiment  field  is  increased.  It  is  obvious 
that  the  substitution  of  3-row  plots  with  discarded  borders  for  single 
rows  will  result  in  greater  variability,  and  will  require  increased  rep- 
lication for  the  same  degree  of  accuracy. 

From  the  foregoing  statements  it  will  be  clear  that  the  number  of 
replications  necessary  for  a given  degree  of  accuracy  may  vary  con- 
siderably with  conditions.  The  number  to  be  used  in  any  specific  ex- 
periment should  be  determined  from  the  variability  of  the  field  in 
question  and  the  degree  of  accuracy  required.  The  variability  of  the 
check  plots  is  usually  considered  a measure  of  the  variability  of  the 
field.  But  when  the  number  of  replications  to  be  used  or  the  extent  of 
experimental  error  is  determined  from  the  variability  of  the  check 
plots,  it  is  assumed  that  the  variability  of  different  varieties  of  the  same 
crop  is  approximately  the  same  under  the  same  conditions.  This  of 
course  is  not  strictly  true.  The  yield  of  two  varieties  may  be  deter- 
mined by  very  different  factors,  as  has  been  stated,  and  their  relative 
variability  may  also  be  quite  different.  The  variability  of  120  plots 


Table  28. — Soil  Heterogeneity  oe  an  Experiment  Field  as  Determined  From 
Yields  oe  Two  Check  Varieties. 

Oats  Variety  and  Strain  Tests.  1921. 


Number 

Average 

Probable 

error  of  a 

Check  variety 

of  plots 

yield 

bu. 

Standard  deviation 
bu.  % 

single  yield  determination 
bu.  % 

Kherson 

120 

37.95 

4.61 

12.15 

3.11 

8.20 

Red  Rustproof 

120 

22.44 

3.99 

17.78 

2.69 

11.99 

each  of  Kherson  and  Red  Rustproof  oats,  grown  side  by  side  as  check 
plots  in  the  oats  variety  and  strain  test  of  1921,  illustrate  the  possibil- 
ity of  a serious  error  in  the  use  of  the  standard  deviation  of  check 
plots  as  a measure  of  the  variability  of  an  experiment  field.  These 
determinations  are  shown  in  Table  28. 

The  field  would  have  been  considered  decidedly  less  variable  if 
Kherson  had  been  used  as  the  check  variety  than  if  Red  Rustproof  had 


54 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


been  used.  Both  of  these  are  standard  recommended  varieties  for  the 
region,  though  they  differ  decidedly  in  their  characteristics.  Both 
have  been  used  frequently  as  check  varieties  at  the  Missouri  station  in 
past  seasons.  From  the  variability  of  the  Kherson  check  plots  the  mean 
yield  of  four  replicate  plots  in  this  experiment  would  be  considered 
to  have  a probable  error  of  4.10  per  cent ; from  the  Red  Rustproof  plots 
the  same  determination  would  be  given  a probable  error  of  6.00  per 
cent.  A degree  of  precision  for  which  we  would  assume  four  replica- 
tions necessary,  judging  from  the  Kherson  check,  would  require  nine 
replications  according  to  the  yields  of  the  Red  Rustproof  check. 

The  importance  of  choosing  a check  variety  typical  of  the  va- 
rieties tested,  if  its  variability  is  to  be  considered  a criterion  of  the 
variability  of  the  field,  is  obvious.  Whether  it  is  possble  to  choose 
a “typical  variety”  for  the  purpose,  in  the  case  of  ordinary  variety 
tests,  remains  to  be  seen. 

ADJUSTMENT  OF  YIELDS  BY  MEANS  OF  CHECK  PLOTS 

Adjustment  of  plot  yields  by  the  use  of  check  plots  has  been  a 
common  practice  in  field  experiments  during  recent  years.  It  is 
recognized  that  no  experiment  field  is  perfectly  uniform  in  produc- 
tivity, and  the  attempt  is  made,  by  means  of  the  check  plot  adjustment, 
to  compensate  the  varieties  or  treatments  which  chance  to  be  located 
on  the  less  productive  plots  for  the  resulting  loss  in  yield.  The  com- 
mon method,  in  variety  tests,  is  to  distribute  over  the  field,  as  fre- 
quently as  practicable,  check  plots  planted  to  the  same  variety  and 
similarly  handled  in  every  way.  The  variation  in  yield  among  these 
check  plots  is  then  considered  a measure  of  the  productivity  of  the 
soil.  By  various  methods,  differing  only  in  detail,  the  yields  of  the 
test  plots  in  parts  of  the  field  giving  high  check  yields  are  reduced,  and 
those  of  test  plots  in  parts  giving  low  check  yields  are  increased,  in 
proportion  to  the  productivity  of  the  soil,  as  indicated  by  the  yields 
of  neighboring  check  plots. 

Previous  Investigation. — Several  investigations  of  the  effect  of 
such  adjustment  on  the  variability  of  replicate  plots  have  been  re- 
ported. The  majority  of  these  have  been  conducted  in  connection  with 
experiments  of  the  type  discussed  in  the  preceding  section,  in  which 
uniformly  handled  fields  have  been  harvested  in  small  sections.  Cer- 
tain of  these  sections,  or  plots,  have  been  considered  check  plots,  and 
on  the  basis  of  their  yields  the  yields  of  the  remaining  plots  have  been 


Experiments  in  Field  Plot  Technic 


55 


adjusted.  The  reduction  of  variability  of  the  adjusted  plot  yields  is 
the  measure  of  the  efficiency  of  the  method. 

Morgan13  reports  an  experiment  of  this  sort,  in  which  63  plots, 
planted  first  to  wheat  and  then  to  fodder  corn,  in  the  same  season, 
were  used.  The  variability  of  the  plot  yields  was  steadily  reduced  as 
the  number  of  check  plots  was  increased. 

In  a similar  experiment  reported  by  Lyon11,  in  which  37  replicate 
1/100  acre  plots  of  corn  were  harvested,  the  use  of  checks  in  every 
second  or  third  plot  was  found  to  reduce  variability,  but  they  were  of 
little  value  when  farther  apart. 

Montgomery14  states  that  alternating  check  plots  with  test  plots 
gives  a high  degree  of  accuracy,  but  the  total  number  of  plots  required 
when  this  method  is  used  is  greater  than  when  the  same  degree  of  ac- 
curacy is  attained  by  the  use  of  replication. 

Kiesselbach5  reports  a comprehensive  trial  of  three  methods  of 
adjusting  yields  by  means  of  check  plots  in  a uniform  field  of  207 
1/30-acre  plots  of  Kherson  oats.  The  effect  on  plot  variability  is  shown 
in  Table  29. 

Table  29. — Effect  on  Plot  Variability  of  Adjusting  Yields  by  Check 
Plots  (Kiesselbach). 

Coefficient 

Method  of  of  variability 

adjustment  Actual  Adjusted 

yields  yields 

Alternate  check  plots. 

Correction  based  on 
average  of  two  ad- 
jacent checks  7.85  7.01 

Checks  every  third  plot. 

Correction  based  on  one 

adjacent  check  plot  7.79  7.35 

Checks  every  third  plot. 

Correction  by  progres- 
sive method,  based  on 

two  nearest  checks  7.87  6.57 


From  these  results  Kiesselbach  concludes  “The  yield  of  system- 
atically distributed  check  plats  cannot  be  regarded  as  a reliable  meas- 
ure for  correcting  and  establishing  correct  theoretical  or  normal 
yields  for  the  intervening  plats.” 

It  should  be  noted  at  this  point  that  even  if  adjustment  by  check 
yields  were  found  invariably  effective  in  experiments  of  this  sort, 


56 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


its  value  in  ordinary  variety  testing  would  not  be  definitely  estab- 
lished. The  practice  involves  not  only  the  assumption  that  the  yields 
of  the  check  plots  are  a fair  indication  of  the  productivity  of  the  in- 
tervening plots  for  the  check  variety,  but  the  further  assumption  that 
different  varieties  respond  similarly  to  differing  growing  conditions. 
Adjustment  of  yields  should  therefore  give  better  results  in  such  ex- 
periments as  those  cited  above  than  it  could  be  expected  to  give  in 
actual  variety  tests. 

This  point  is  well  illustrated  by  observations  reported  by  Salmon18. 
Two  varieties  of  barley,  Gatami  and  Odessa,  were  grown  side  by  side 
in  fiftieth-acre  plots  in  five  distributed  portions  of  a field.  Gatami  gave 
an  average  yield  of  18.3  bushels  per  acre,  with  quite  uniform  yields  in 
the  five  plots,  as  evidenced  by  their  probable  error  of  0.68  bushel, 
while  Odessa  yielded  13.3  bushels  per  acre  in  the  first  plot,  6.35  bushels 
per  acre  in  the  second,  and  a negligible  yield  in  the  other  three.  Ob- 
viously the  adjustment  of  the  yield  of  either  of  these  varieties  on  the 
basis  of  the  other  variety  as  a check,  would  enormously  increase 
rather  than  decrease  the  experimental  error.  As  Salmon  points  out, 
an  error  similar  in  kind  though  less  in  degree  may  occur  commonly 
in  variety  tests,  when  the  yields  of  varieties  are  determined  by  dif- 
ferent limiting  factors.  And  if  this  is  generally  the  case,  adjustment 
by  check  yields  will  be  of  doubtful  value,  even  if  it  were  found  to 
eliminate  variability  completely  in  uniform  plot  tests. 

There  is  a growing  tendency,  consequently,  to  discontinue  the  use 
of  check  plots  for  adjusting  yields  in  variety  tests,  and  to  use  them 
only  to  measure  soil  variability  and  to  indicate  the  degree  of  error 
in  yield  determinations  of  the  tested  varieties.  Adjustment  of  yields 
has  never  been  as  common  in  preliminary  tests  as  in  tests  on  larger 
plots,  principally  because  of  the  great  amount  of  computation  neces- 
sary in  adjusting  the  yields  of  ten  or  twenty  replicate  rod-rows  of  a 
large  number  of  varieties,  and  because  the  yield  of  a single  rod-row, 
exposed  to  varying  competition  and  materially  affected  by  small  me- 
chanical errors,  is  at  best  a very  unreliable  measure  of  productivity  on 
which  to  base  the  adjustment  of  the  yields  of  several  other  plots. 

Experimental  Results. — It  would  of  course  be  very  desirable  to 
use  check  plots  for  reducing  plot  variability,  if  the  method  could  be 
relied  on,  because  of  the  economy  of  the  practice.  The  only  certain 
method  of  reducing  plot  variability  is  by  means  of  replication,  and  it 
may  be  considered  a fairly  general  rule  that  the  variability  of  plots 
on  a given  field,  as  measured  by  the  standard  deviation  or  the  prob- 
able error,  will  in  general  be  reduced  by  replication  in  proportion  to 
the  square  root  of  the  number  of  replications.  In  other  words,  the 


Experiments  in  Fieed  Plot  Technic 


57 


variability  of  the  mean  of  16  replicate  plots  will  be  about  half  that  of 
the  mean  of  4 replicate  plots.  Now  the  maximum  use  of  check  plots, 
that  is,  the  practice  of  alternating  check  plots  and  test  plots,  requires 
the  same  land  and  labor  as  would  be  required  by  doubling  the  num- 
ber of  replications,  if  no  check  plots  were  used.  As  doubling  the 
number  of  replications  will  in  general  give  a standard  deviation  about 
equal  to  the  original  standard  deviation  divided  by  the  square  root  of 

2,  it  will  reduce  variability  about  30  % = -7071  ^ . If  alternat- 

ing with  check  plots  will  consistently  reduce  variability  more  than  30 
per  cent  it  will  be  generally  a more  economical  way  to  control  error. 
Similarly,  the  use  of  check  plots  in  every  third  plot  requires  as  much 
land  as  would  be  required  by  increasing  the  number  of  replications  by 
50  per  cent  (using  three  replications  instead  of  two,  or  fifteen  instead 
of  ten).  From  this  relation  the  reduction  of  variability  necessary  if 
this  practice  is  to  equal  replication  in  effectiveness  can  be  easily  com- 
puted. Such  determinations  for  check  plots  at  various  intervals  are 
shown  in  Table  30. 


Table  30. — Reduction  of  Variability  by  the  Use  of  Check  Plots  Equivalent 
to  That  Probably  Attainable  With  the  Same  Number 
of  Plots  by  Replication. 


Distribution  of 
check  plots 

Equivalent  increase  in 
number  of  replications 
% 

Reduction  in 
standard  deviation  to 
be  expected  by  such 
increase  in  replication 
% 

Alternate  plots 

100.00 

29.29 

Every  third  plot 

50.00 

18.35 

Every  fourth  plot 

33.33 

13.50 

Every  fifth  plot 

25.00 

10.55 

Every  sixth  plot 

20.00 

8.71 

Every  seventh  plot 

16.67 

7.41 

Every  eighth  plot 

14.29 

6.47 

Every  ninth  plot 

12.50 

5.75 

Every  tenth  plot 

11.11 

5.12 

If  protected  single-row  or  3-row  plots  are  used  in  preliminary 
experiments  a more  reliable  measure  of  soil  productivity  is  available, 
and  consequently  the  adjustment  of  yields  is  more  likely  to  be  of  value, 
than  when  unprotected  single-row  plots  are  used.  By  the  use  of 
planting  plans  of  the  sort  employed  in  these  experiments,  it  is  pos- 


58  Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 

sible  to  adjust  the  yields  by  a somewhat  shortened  method.  If  adjust- 
ment of  yield  is  effective  in  reducing  plot  variability  in  this  sort  of 
test  it  can  be  accomplished  with  but  little  increase  in  labor.  In  each 
of  the  tests  reported  in  this  paper  a trial  of  the  effectiveness  of  adjust- 
ing yields  by  means  of  check  plots  was  made,  the  criterion  of  accuracy 
being  in  each  case  the  variability  of  the  yields  of  the  replicate  plots  of 
each  variety.  Since  the  number  of  replicate  plots  was  only  three 
or  four  the  average  deviation  was  determined  instead  of  the  stand- 
ard deviation. 

Method  Used  in  Adjusting  Yields. — The  method  employed  in  ad- 
justing yields  may  be  described  as  follows:  The  average  yield  of  all 
check  plots  and  the  relative  yield  of  each  check  plot  in  terms  of  this 
average  (that  is,  the  quotient  obtained  by  dividing  the  yield  of  the  in- 
dividual check  plot  by  the  average  yield  of  all  check  plots)  were  de- 
termined. The  relative  yield  of  each  check  plot,  expressed  in  per- 
centage of  the  mean  check  yield,  is  designated  hereafter  as  the  “plot 
value”  of  that  check  plot.  When  the  average  yield  of  all  check  plots 
is  25  bushels  per  acre,  the  plot  value  of  a check  plot  yielding  30  bushels 
per  acre  is  120  per  cent — in  other  words  it  is  20  per  cent  more  pro- 
ductive than  the  average.  Now  assuming  gradual  change  in  the  pro- 
ductivity of  the  soil  between  check  plots,  each  test  plot  is  assigned  a 
plot  value  by  interpolation.  The  adjusted  yield  of  each  plot  is  then 
determined  by  dividing  the  actual  yield  by  the  plot  value. 

The  short  method  for  adjusting  yields,  referred  to  above,  is 
based  on  the  fact  that  the  varieties  occur  in  the  same  order  in  each 
series.  Thus  in  the  field  diagrammed  in  figure  1,  the  following  se- 
quence of  plots  occurs  in  each  of  the  four  series : 

ck  1 17  33  49  65  81  ck 

Now  if  the  average  yield  of  the  four  check  plots  adjoining  variety 
1,  and  the  average  yield  of  the  four  check  plots  adjoining  variety  81 
are  each  given  a plot  value,  corresponding  plot  values  for  the  mean 
yields  of  varieties  1,  17,  33,  49,  65,  and  81  may  be  interpolated,  and 
the  mean  yields  may  be  adjusted  in  one  operation.  The  same  method 
may  be  used,  of  course,  regardless  of  the  number  of  replications.  The 
result  will  not  be  exactly  the  same  as  that  of  averaging  the  adjusted 
yields  determined  individually,  but  will  in  most  cases  approximate  it 
closely,  the  slight  difference  being  caused  by  the  disproportion  of  yield 
and  plot  value  in  the  plots  averaged.  It  is  doubtful  that  either  meth- 
od is  consistently  more  accurate  than  the  other. 

When  the  check  plot  yield  is  used  in  the  adjustment  of  the  yields 
of  other  plots  it  is  of  course  essential  that  it  should  be  a reliable  de- 
termination, not  unduly  affected  by  factors  not  affecting  the  neighbor- 


Experiments  in  Field  Plot  Technic 


59 


ing  plots.  For  example  if  the  yield  of  a check  plot  is  reduced  20  per 
cent  by  a poor  stand,  the  adjusted  yields  of  neighboring  plots  will  be 
increased  to  the  same  extent  as  if  the  check  plot  yield  had  been  low  be- 
cause of  poor  soil,  and  will  consequently  be  considerably  higher  than 
they  should  be.  It  is  important  therefore  that  conditions  be  made  as  fa- 
vorable as  possible  for  accurate  yield  testing  when  this  method  is  used. 
One  cause  for  poor  results  in  the  adjustment  of  yield  in  some  of  the 
experiments  reported  in  this  paper  was  failure  to  protect  the  outside 
strip  of  check  plots  by  means  of  border  rows,  in  a few  of  the  tests,  be- 


Table  31. — Relative  Variability  of  Actual  and  Adjusted  Yields. 
Average  Deviation  in  Percentage  of  Yield. — Barley  Variety  Test  1919. 


Planting 

number 

Variety  3 

Average  deviation 
Actual  yields  Adj  usted 

interior  rows  5 rows  3 interior  rows 
% % % 

i yields 
5 rows 
% 

1 

Hanna  906 

19.81 

17.82 

13.15 

10.80 

2 

Steigum  907 

15.17 

18.79 

13.48 

8.40 

3 

Luth  908 

29.97 

28.17 

5.51 

4.62 

4 

Eagle  913 

26.14 

29.79 

9.71 

12.98 

6 

Servian  915 

18.37 

17.97 

7.36 

5.59 

7 

Odessa  916 

2.31 

4.33 

23.99 

17.01 

8 

Lion  923 

14.65 

12.10 

11.37  . 

11.03 

10 

Horn  926 

4.08 

2.00 

16.45 

12.74 

11 

Odessa  927 

13.62 

9.16 

21.62 

13.76 

12 

Summit  929 

5.28 

6.45 

14.23 

11.59 

13 

Mariout  932 

11.02 

11.57 

18.68 

14.31 

14 

Odessa  934 

13.73 

13.91 

11.18 

10.31 

15 

Peruvian  935 

13.25 

17.87 

12.42 

16.82 

16 

Trebi  936 

11.27 

12.53 

18.78 

18.28 

18 

Oderbrucker  940 

10.77 

14.46 

13.60 

15.98 

19 

Frankish  953 

20.53 

19.65 

22.63 

18.49 

20 

Manchuria  956 

6.88 

6.33 

13.89 

10.68 

21 

Oderbrucker  957 

17.88 

13.62 

1.97 

3.27 

22 

Manchuria  x Champi 
of  Vermont 

on  39.47 

39.19 

21.93 

20.35 

23 

Luth  972 

16.77 

18.61 

7.05 

7.48 

24 

Red  River  973 

13.94 

11.02 

12.37 

12.33 

25 

Featherston  1118 

21.40 

20.89 

8.47 

13.39 

26 

Featherston  1119 

16.59 

15.25 

4.78 

12.26 

27 

Featherston  1120 

15.91 

13.64 

2.48 

6.44 

28 

Hanna  x Champion 
of  Vermont  1121 

16.00 

16.81 

28.36 

28.50 

29 

Manchuria  1125 

6.79 

14.42 

7.78 

1.55 

30 

Malting  1129 

12.86 

10.40 

5.40 

9.02 

Mean 

15.35 

15.44 

12.91 

12.15 

60 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


cause  of  lack  of  space.  The  check  plots  growing  on  the  border  of  the 
field  were  materially  reduced  in  yield,  in  some  cases,  notably  the  oats 
strain  test  of  1919  and  the  wheat  variety  test  of  1921.  In  these  cases 
the  variability  of  the  actual  and  adjusted  yields  has  been  computed 
both  for  all  series  and  for  the  remaining  series  when  the  one  affected 
by  an  unreliable  check  is  discarded. 

Relative  Variability  of  Actual  and  Adjusted  Yields. — The  relative 
variability  of  actual  and  adjusted  yields  of  both  3-row  and  5-row  plots 
in  the  barley  variety  test  is  shown  in  Table  31.  In  this  test  there  were 
three  replications,  and  the  check  variety  was  Oderbrucker,  in  every 


Table  32. — Relative  Variability  oe  Actual  and  Adjusted  Yields. 
Average  Deviation  in  Percentage  of  Yield 
Oats  Variety  Test  1919. 


Planting 

number  Variety 

Average  Deviation 
3 Series  4 Series 

(3  interior  rows)  (3  interior  rows) 

Actual  Adjusted  Actual  Adjusted 

yields  yields  yields  yields 

% % % % 

1 

A.  sterilis  nigra 

4.32 

1.46 

9.18 

5.02 

2 

Black  Mesdag 

9.03 

9.69 

7.29 

12.90 

3 

C.  I.  602 

13.72 

16.00 

16.30 

13.63 

3 

C.  I.  603 

4.72 

3.09 

5.84 

3.88 

5 

C.  I.  620 

4.73 

10.14 

11.24 

13.67 

6 

Early  Champion 

18.63 

15.18 

14.97 

14.65 

7 

Early  Gothland 

14.20 

4.67 

11.55 

4.18 

8 

Garton  473 

5.99 

6.25 

8.42 

9.42 

9 

Garton  585 

14.08 

19.44 

16.92 

19.95 

10 

Golden  Giant 

9.44 

14.31 

14.40 

15.09 

11 

Irish  Victor 

9.69 

3.29 

7.72 

16.40 

12 

Japanese  Selection 

6.87 

4.71 

11.85 

5.20 

13 

June 

18.37 

11.19 

17.53 

10.37 

14 

Kherson  Selection 

17.01 

9.20 

15.06 

20.36 

15 

Fulghum 

9.69 

11.36 

13.06 

17.32 

16 

Lincoln 

21.07 

12.54 

16.56 

11.83 

17 

Monarch 

6.12 

4.55 

9.06 

33.36 

18 

North  Finnish 

8.69 

5.17 

7.84 

27.03 

19 

Scottish  Chief 

5.05 

4.28 

5.10 

15.42 

20 

Sparrow  bill  (Missouri) 

10.98 

10.82 

12.38 

13.15 

21 

Sparrow  bill  (Cornell) 

4.45 

3.25 

12.11 

3.85 

22 

Tobolsk  1 

6.17 

3.85 

13.92 

5.38 

23 

Tobolsk  2 

11.56 

9.24 

20.35 

13.96 

24 

White  Tartar 

10.94 

4.75 

9.51 

4.54 

Mean 

10.23 

8.27 

12.01 

12.94 

Experiments  in  Field  Plot  Technic 


61 


sixth  plot.  As  a result  of  the  adjustment  of  yields,  the  average  devia- 
tion of  3-row  plots  was  reduced  from  15.35  per  cent  to  12.91  per  cent, 
a reduction  of  16  per  cent,  and  that  of  5-row  plots  from  15.44  per 
cent  to  12.15  per  cent,  a reduction  of  21  per  cent. 

The  relative  variability  of  actual  and  adjusted  yields  in  the  oats 
variety  test  of  1919  is  shown  in  Table  32.  In  this  field  the  check, 
Red  Rustproof,  was  in  every  ninth  plot.  When  the  series  affected  by 
the  faulty  check  yields  of  the  border  plots  is  included  the  variability  of 
the  adjusted  yields  is  slightly  higher  than  that  of  the  actual  yields, 
but  when  this  series  is  discarded  the  average  variability  as  measured 
by  the  mean  deviation  is  reduced  19  per  cent. 

It  might  be  expected  that  the  oats  strains  grown  on  the  same  field 
would  show  a greater  reduction  of  variability  than  the  varieties,  since 
practically  all  of  them  were  of  the  same  variety  as  the  check,  and  since 


Table  33. — Relative  Variability  oe  Actual  and  Adjusted  Yields. 
Average  Deviation  in  Percentage  of  Yield. 

Oats  Strains  Test  1919. 


Planting 

number 

Accession 

number 

Average  deviation 

Actual  yields  Adjusted  yields 

3 interior  rows  5 rows  3 interior  rows  5 rows 
% % % % 

1 

0119 

10.30 

7.75 

11.22 

8.81 

2 

0120 

4.70 

6.58 

3.01 

1.76 

3 

0121* 

5.76 

3.25 

4.57 

4.14 

4 

0122 

4.62 

3.52 

6.46 

3.82 

5 

0123 

9.91 

8.25 

11.47 

9.62 

6 

0125 

3.18 

3.34 

5.39 

6.60 

7 

0126 

7.62 

5.95 

10.56 

16.58 

8 

0127* 

6.76 

5.09 

6.92 

9.85 

9 

0124* 

6.13 

6.34 

4.92 

4.59 

10 

0133 

7.07 

4.77 

3.92 

5.36 

11 

0128 

4.17 

3.56 

3.58 

3.36 

12 

0129 

5.02 

7.07 

5.94 

6.35 

13 

0130 

4.20 

2.62 

6.74 

9.72 

14 

0131 

2.59 

2.59 

4.98 

2.94 

15 

0132 

7.38 

5.81 

12.38 

12.08 

Mean 

5.96 

5.10 

6.80 

7.04 

* Not  taxonomically  Red  Rustproof. 


the  check  plots  were  more  frequent,  being  in  every  sixth  plot.  The 
results  of  adjusting  yields  in  this  test,  both  for  protected  3-row  plots 
and  for  unprotected  5-row  plots  in  four  series  are  shown  in  Table  33. 
Contrary  to  expectation,  the  variability  was  not  reduced  by  adjustment 


Table  34. — Relative  Variability  oe  Actual  and  Adjusted  Yields. 

Average  Deviation  in  Percentage  of  Yield. — Wheat  Variety  Test  1920. 


62 


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Experiments  in  Fieed  Plot  Technic 


63 


of  yield.  A possible  explanation  is  the  extremely  low  variability  of  the 
actual  yields,  indicating  that  the  field,  which  was  quite  small,  was  re- 
latively uniform.  Any  gain  in  uniformity  from  a check  adjustment 
of  yields  would  of  course  be  expected  to  be  greater  in  highly  variable 
than  in  more  uniform  fields.  The  relative  uniformity  of  this  field 
is  indicated  not  only  by  the  low  mean  deviation  of  the  test  plots,  but 
also  by  the  low  standard  deviation  of  the  check  plots,  which  was  only 
11.68  per  cent,  as  compared  with  a standard  deviation  of  22.59  per 
cent  in  the  check  plots  of  the  adjoining  oats  variety  test. 

The  effect  of  adjusting  yields  on  the  variability  of  3-row  and 
5-row  plots  in  the  wheat  variety  test  of  1920  is  shown  in  Table  34. 
In  this  test  the  check  variety,  Fultz,  was  grown  in  every  seventh  plot. 
There  were  four  series  of  the  ninety-six  varieties. 

The  reduction  in  variability  was  very  marked,  being  37  per  cent 
for  3-row  plots  and  42  per  cent  for  5-row  plots.  The  variability  of 
almost  every  variety  was  reduced,  and  the  reliability  of  the  results 
was  undoubtedly  much  increased. 

The  wheat  variety  test  of  1921,  occupying  an  equal  area  on  a 
neighboring  field,  and  with  similar  varieties  and  the  same  planting 
plan,  gave  decidedly  different  results.  In  this  field  the  check  va- 
riety was  Poole.  Several  check  plots  on  the  border  were  abnormal, 
and  the  computations  are  therefore  given  both  for  three  series  and  for 
four,  the  series  affected  by  the  abnormal  check  yields  being  dis- 
carded in  the  former  case.  The  relative  variability  of  actual  and  ad- 
justed yields  is  shown  in  Table  35. 

Although  the  check  yields  are  somewhat  less  variable  for  three 
series  than  for  four,  the  adjustment  was  not  effective  in  either  case  in 
reducing  variability.  The  adjusted  yields  are  10  per  cent  more  va- 
riable than  the  actual  yields  for  the  three  series  and  34  per  cent  higher 
for  the  four. 

Similar  results  were  obtained  in  the  wheat  mixture  test  of  the 
same  season,  in  which  several  of  the  same  varieties  were  included, 
and  the  same  check  variety  was  used.  In  this  test  the  check  variety 
was  in  every  sixth  plot,  and  four  replications  were  used.  The  results 
of  adjusting  yields  are  shown  in  Table  36.  Variability  was  increased 
from  9.84  per  cent  to  13.81  per  cent,  an  increase  of  40  per  cent.  Thus 
the  results  of  adjusting  yields  of  wheat  varieties  in  1921  are  directly 
contrary  to  the  results  of  the  same  practice  in  1920. 

Difference  in  Results  Obtained  by  Adjustment  with  Different 
Check  Varieties. — In  the  oats  variety  and  strain  tests  of  1921,  two 
check  varieties,  Kherson  and  Red  Rustproof,  were  grown.  In  these 
tests  96  strains  were  included,  32  of  Kherson,  32  of  Red  Rustproof,  and 


Table  35. — Relative  Variability  of  Actual  and  Adjusted  Yields. 
Average  Deviation  in  Percentage  of  Yield. — Wheat  Variety  Test  1921, 


64 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


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Experiments  in  Field  Plot  Technic 


65 


Table.  36 — Relative  Variability  of  Actual  and  Adjusted  Yields. 
Average  Deviation  in  Percentage  of  Yield.  Wheat  Mixture  Test  1921 


Planting 

number  Variety 

Average  Deviation 
Actual  yields  Adjusted  yields 

(3  interior  (3  interior 

rows)  rows) 

% % 

1 

Fulcaster 

9.67 

16.90 

2 

Harvest  Queen 

9.35 

21.47 

3 

Mixture  No.  1 

6.12 

20.11 

4 

Michigan  Wonder 

7.88 

20.51 

5 

Nigger 

2.33 

20.16 

6 

Michigan  Wonder  No.  31 

4.93 

13.86 

7 

Michigan  Wonder  No.  54 

12.96 

16.03 

8 

Mixture  No.  2 

15.38 

17.86 

9 

Michigan  Wonder  No.  96 

4.99 

13.88 

10 

Michigan  Wonder  No.  209 

5.30 

14.83 

11 

Beechwood  Hybrid  No.  12 

9.04 

11.01 

12 

Beechwood  Hybrid  No.  85 

14.98 

15.49 

13 

Mixture  No.  3. 

16.76 

10.47 

14 

Beechwood  Hybrid  No.  87 

10.16 

11.94 

15 

Beechwood  Hybrid  No.  207 

20.80 

11.81 

16 

Michigan  Wonder  No.  221 

7.13 

8.90 

17 

Kanred 

10.09 

12.86 

18 

Mixture  No.  4 

10.39 

8.44 

19 

New  York  123-32 

16.73 

12.09 

20 

Red  Rock 

14.04 

10.68 

21 

Red  Hussar 

12.71 

17.42 

22 

Turkey  (Kansas) 

17.32 

13.74 

23 

Mixture  No.  5 

2.87 

12.73 

24 

Michigan  Amber 

3.39 

10.71 

25 

Nigger 

4.09 

10.47 

26 

Fulcaster  (Co-op) 

2.09 

14.25 

27 

Fulcaster  (Outl) 

11.97 

17.18 

28 

Mixture  No.  6 

11.19 

7.75 

29 

Fulcaster  (Blazier) 

11.41 

6.77 

30 

Fulcaster  (Cowles) 

9.09 

14.11 

Mean 

9.84 

13.81 

32  of  other  varieties.  The  yields  were  adjusted  by  means  of  each 
check  variety  separately,  to  determine  the  relation  between  the  ef- 
fectiveness of  yield  adjustment  and  the  similarity  of  the  check  to  the 
tested  variety.  The  results  of  this  adjustment  on  plot  variability  are 
shown  in  Tables  37  and  38. 

The  variability  of  the  yields  of  the  Red  Rustproof  strains  was 
somewhat  reduced  (6  per  cent)  by  adjustment  according  to  the  yields 


66 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


Table  37. — Relative  Variability  of  Actual  and  Adjusted  Yields. 
Average  Deviation  in  Percentage  of  Yield.  Oats  Variety  Test  1921 


Planting 

number  Variety 

Average  deviation 

Actual  yields  Adjusted  yields 

(3  interior  (3  interior  rows) 

rows)  (Kherson)  (Red  Rustproof) 

% % % 

65 

Burt 

10.22 

11.54 

11.00 

66 

Canadian 

9.29 

8.93 

18.37 

67 

C.  I.  603 

9.21 

4.94 

13.89 

68 

Culberson 

6.46 

8.80 

4.44 

69 

Danish  Island 

10.53 

8.83 

12.56 

70 

Early  Dakota 

9.33 

7.96 

11.45 

71 

Early  Gothland 

10.00 

19.16 

14.69 

72 

Garton  748 

14.35 

13.26 

14.77 

73 

Green  Russian 

9.75 

11.23 

13.75 

74 

Irish  Victor 

9.43 

8.07 

15.99 

75 

Joanette 

30.75 

29.36 

26.94 

76 

Fulghum  042 

5.19 

8.50 

8.99 

77 

Monarch 

4.76 

6.36 

8.54 

78 

Monarch  Selection 

2.89 

5.74 

11.55 

79 

Scottish  Chief 

10.95 

15.01 

8.66- 

80 

Silvermine  050 

11.24 

9.88 

9.21 

81 

Silvermine  Selection 

11.25 

16.14 

3.16 

82 

Sparrowbill  (C) 

23.93 

23.03 

25.23 

83 

Sterilis  Selection 

8.02 

7.30 

9.17 

84 

Storm  King 

11.68 

13.94 

13.15 

85 

Swedish  Select  057 

14.48 

14.89 

11.80 

86 

Fulghum  065 

8.81 

11.04 

9.41 

87 

Fulghum  0113 

22.19 

11.18 

15.85 

88 

Silvermine  0115 

8.38 

6.29 

3.59 

89 

Silvermine  0117 

6.90 

8.95 

9.69 

90 

Fulghum  0124 

5.17 

3.38 

7.89 

91 

Fulghum  0145 

9.81 

8.63 

15.66 

92 

Fulghum  0149 

10.19 

10.77 

15.70 

93 

Fulghum  0151 

10.85 

12.71 

10.80 

94 

Fulghum  0152 

9.07 

14.59 

19.61 

95 

Silvermine  0165 

5.63 

11.32 

4.62 

96 

Swedish  Select  0165 

16.74 

14.02 

8.74 

Mean 

10.86 

11.43 

12.15 

of  the  Red  Rustproof  check,  but  was  slightly  increased  (2  per  cent) 
when  the  Kherson  check  was  used.  On  the  other  hand,  the  variability 
of  the  yields  of  the  Kherson  strains,  though  not  reduced  by  either  check, 
was  increased  only  4 per  cent  by  the  Kherson  check,  while  it  was  in- 
creased 48  per  cent  by  the  Red  Rustproof  check.  Neither  check  was 
effective  in  adjusting  the  yields  of  the  other  varieties,  the  Kherson 


Experiments  in  Field  Plot  Technic 


67 


Tabu:  38. — Relative  Variability  oe  Actual  and  Adjusted  Yields. 
Average  Deviation  in  Percentage  of  Yield. — Oats  Strain  Test  1921 
(Red  Rustproof  and  Kherson) 


Red  Rustproof  strains 


Kherson  strains 


Planting  Number....  | 

Strain 

Average  deviation 

Planting  Number., 

Strain 

Average  deviation 

Actual  yields  (3 
interior  rows) 

Adjusted  yields 
(3  interior  rows) 

Actual  yields  (3  in- 
terior rows) 

Adjusted  yields 
(3  interior  rows) 

(Kher-I 
son) . . 1 

(Red 
Rust 
proof ) 

(Kher- 
son) . . 

(Red 

Rust- 

proof) 

% 

% 

% 

% 

% 

% 

1 

066 

18.13 

15.48 

13,04 

2 

023 

15.80 

7.59 

12.93 

3 

067 

12.89 

14.19 

11.36 

4 

040 

3.86 

5.59 

10.45 

5 

068 

23.14 

21.34 

12.47 

6 

041 

9.50 

5.82 

4.07 

7 

069 

21.55 

20.54 

11.70 

8 

052 

3.46 

2.83 

10.49 

9 

072 

14.29 

12.26 

10.34 

10 

053 

6.26 

5.97 

10.14 

11 

074 

12.18 

17.23 

13.94 

12 

079 

11.49 

9.91 

10.20 

13 

075 

13.40 

18.28 

18.80 

14 

080 

1.72 

7.02 

13.20 

15 

0118 

28.77 

32.06 

26.92 

16 

082 

6.19 

8.74 

15.01 

18 

0119 

10.32 

13.47 

13.62 

17 

083 

5.31 

5.88 

9.26 

20 

0120 

7.28 

6.11 

12.32 

19 

085 

8.69 

10.65 

7.30 

22 

0122 

17.16 

12.32 

14.20 

21 

086 

11.73 

7.45 

8.45 

24 

0125 

10.91 

11.52 

8.40 

23  Mixture***  4.87 

3.84 

9.19 

26 

0126 

10.41 

10.24 

4.66 

25 

088** 

13.04 

13.79 

9.16 

28 

0128 

14.48 

17.36 

2.76 

27 

089 

4.69 

6.77 

12.89 

30 

0129 

7.89 

6.31 

7.44 

29 

090 

2.84 

4.85 

10.64 

32 

0130 

7.99 

11.78 

9.17 

31 

091 

12.46 

14.67 

10.84 

33 

0131 

12.43 

13.96 

13.23 

34 

094 

4.59 

6.43 

6.57 

35 

0132 

17.77 

14.67 

12.11 

36 

095 

4.36 

4.96 

14.62 

37 

0133 

14.07 

11.63 

13.55 

38 

096 

4.39 

7.71 

4.66 

39 

0134 

29.91 

30.37 

29.50 

40 

097 

5.53 

6.97 

11.80 

41 

0135 

14.36 

17.58 

13.11 

42 

098 

9.09 

10.03 

13.18 

43 

0136* 

7.80 

7.59 

12.69 

44 

099 

6.89 

6.26 

10.20 

45 

0141 

12.58 

13.30 

11.11 

46 

0100 

6.90 

4.65 

8.55 

47 

0163 

2.70 

1.82 

14.28 

48 

0155 

12.75 

4.49 

23.56 

50 

0169 

21.95 

17.79 

23.03 

49 

0157 

6.34 

7.40 

14.18 

52 

0181 

9.95 

7.62 

9.69 

51 

0158 

10.81 

12.24 

6.26 

54 

0182 

26.12 

21.76 

20.84. 

53 

0159 

6.33 

10.82 

8.26 

56 

0183* 

4.52 

5.51 

8.99 

55 

0160 

4.98 

6.12 

12.52 

58 

0383 

10.10 

15.60 

20.18 

57 

0161 

5.55 

9.19 

9.27 

60 

0391 

9.55 

11,37 

6.47 

59 

0162 

11.28 

13.38 

9.47 

62 

0394 

13.03 

11.52 

10.29 

61 

0167 

2.65 

3.37 

4.41 

64 

0395 

8.69 

11.40 

17.96 

63 

0174 

10.74 

8.96 

15.67 

Mean 

14.47 

14.70 

. 13.55 

Mean 

7.16 

7.44 

10.59 

* Not 

taxonomically  Red 

Rustproof. 

Excluded  from  : 

average. 

**  Not  taxonomically  Kherson.  Excluded  from  average. 


•••Mixture  of  strains  082,  094,  0100,  0174. 


check  increasing  their  variability  7 per  cent,  and  the  Red  Rustproof 
check  20  per  cent.  These  results  indicate  the  importance  of  using 
a check  variety  typical  of  the  varieties  tested,  when  adjustment  of 
yields  is  to  be  made ; and  the  danger  of  increasing  rather  than  decreas- 


68 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


ing  error  by  this  practice  when  the  tested  varieties  are  quite  different 
in  habit  from  the  check  variety. 

The  use  of  an  unsuitable  check  variety  not  only  increases  the 
margin  of  error,  but  it  may  cause  very  deceptive  comparative  results. 
For  example,  the  average  yields  of  the  Kherson  strains  0155  and  0157, 
unadjusted  and  adjusted  according  to  the  yields  of  both  check  va- 
rieties, are  shown  below: 


Method 

Strain 

0155 

0157 

Yield 

Average 

Deviation 

Yield 

Average 

Deviation 

Unadjusted 

37.50 

12.75 

43.69 

6.34 

Adjusted  by  Kherson  check 

34.50 

4.49 

43.69 

7.40 

Adjusted  by  Red  Rustproof  check 

39.94 

23.56 

39.38 

14.18 

The  17  per  cent  advantage  in  yield  of  strain  0157  is  increased  to 
27  per  cent  by  the  Kherson  adjustment,  and  since  the  variability  of 
the  replicate  yields  is  reduced  by  the  adjustment  we  may  fairly  as- 
sume that  the  latter  is  the  more  reliable  figure.  But  when  the  Red 
Rustproof  check  is  used  for  adjusting  yields,  the  advantage  of  strain 
0157  disappears  entirely.  The  inaccuracy  of  the  yields  adjusted  by 
Red  Rustproof  is  indicated  by  the  increase  in  plot  variability  result- 
ing from  this  adjustment.  Thus  the  adjustment  of  yields  by  means 
of  check  plots  may  mask  considerable  differences  in  yields  between  the 
varieties  under  test. 

Although  Kherson  and  Red  Rustproof  are  decidedly  different  in 
type,  both  are  commonly  grown  in  Missouri,  and  both  have  been  used 
frequently  here  as  check  varieties  in  oats  variety  tests.  It  is  interesting 


Table  39. — Relative  Variability  of  Actual  and  Adjusted  Yields  of  Kherson 
and  Red  Rustproof  Oats,  Each  in  120  Distributed  Plots. 

Oats  Variety  and  Strain  Tests  1921. 


Standard  deviation 

Yield 

Actual 

Adjusted 

Variety 

Actual  Adjusted 

yield 

yield 

% 

% 

Kherson 

37.95  39.04 

12.15 

20.79 

Red  Rustproof 

22.44  22.80 

17.78 

19.92 

Experiments  in  Field  Plot  Technic 


69 


to  determine  the  effect  on  variability  of  adjusting  the  yields  of  the 
120  plots  of  Kherson,  on  the  basis  of  those  of  the  120  plots  of  Red 
Rustproof  adjoining  them,  and  those  of  the  120  plots  of  Red  Rustproof, 
on  the  basis  of  the  yields  of  the  adjoining  Kherson  plots.  In  this 
adjustment  the  yield  of  each  plot  is  divided  by  the  plot  value  of  the 
adjoining  plot,  and  the  method  corresponds  to  method  II  used  by 
Kiesselbach  in  the  experiment  cited  above  (see  Table  29).  The  results 
of  the  yield  adjustment  are  shown  in  Table  39. 

The  adjustment  of  plot  yields  by  means  of  check  plots  of  a va- 
riety distinctly  different  in  type  resulted  in  a decided  increase  in  plot 
variability,  even  though  the  plot  values  used  were  determined  in  each 
case  by  the  yield  of  the  immediately  adjacent  plot.  If  the  yields 
of  the  Kherson  and  the  Red  Rustproof  plots  had  been  perfectly  ac- 
curate measures  of  the  productivity  of  the  soil,  the  plot  values  of  the 
adjacent  plots  would  have  been  almost  the  same  in  each  of  the  120 
locations,  and  the  adjustment  of  the  yields  of  either  variety  by  those 
of  the  other  would  have  reduced  variability  almost  to  zero.  Instead, 
variability  was  actually  and  very  decidedly  increased,  because  the  sec- 
tions of  the  field  which  gave  relatively  high  yields  of  Kherson,  gave 
relatively  low  yields  of  Red  Rustproof,  and  vice  versa,  in  many  cases. 
In  fact,  there  was  very  little  relation  between  the  productivity  of  a 
portion  of  the  field  as  determined  by  a Kherson  check,  and  the  pro- 


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o 

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80 

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6 

90 

to 

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2 

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10 

100 

to 

110 

1 

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2 

4 

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110 

to 

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4 

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to 

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to 

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140 

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150 

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Total 

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20 

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19 

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120 

Figure  7— Correlation  Between  Yields  of  Kherson  Check  Plots  and 

Yields  of  Adjacent  Red  Rustproof  Check  Plots,  in  Oats  Variety  and 
Strain  Tests  1921. 

r=  +.162  ± .060. 


70 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


ductivity  of  the  same  portion  of  the  field  as  determined  by  an  adjacent 
Red  Rustproof  check  plot.  This  correlation  is  shown  in  figure  7.  The 
coefficient  of  correlation  is  less  than  three  times  its  probable  error — 
the  correlation  has  not  even  statistical  significance ! The  relative  pro- 
ductivity of  different  portions  of  the  field,  as  indicated  by  the  two  check 
varieties,  is  shown  in  figure  8.  If  Kherson  had  been  used  as  a check 


Figure  8. — Relative  Variability  of  Different  Parts  of  an  Experiment 
Field,  as  indicated  by  the  Yields  of  Adjacent  Check  Plots  of  Kherson 
and  Red  Rustproof  Oats.  Oats  Variety  and  Strain  Tests  1921.  In  the 
diagram  on  the  left,  points  of  equal  productivity,  as  indicated  by  the  yields 
of  the  Kherson  check  plots,  are  connected  by  lines  (as  points  of  equal  elevation 
are  connected  by  lines  on  a contour  map).  In  the  diagram  on  the  right,  the 
same  field  is  similarly  mapped  according  to  the  yields  of  the  Red  Rustproof 
check  plots.  The  numbers  indicate  the  plot  values  of  the  points  concerned. 

variety  for  adjusting  yields,  the  yields  of  certain  plots  would  have  been 
increased  to  compensate  for  the  low  productivity  of  the  soil ; if  Red 
Rustproof  had  been  used  the  yields  of  the  same  plots  would  have 
been  decreased  to  compensate  for  the  high  productivity  of  the  same 
soil.  The  fact  is  that  certain  parts  of  the  field  were  actually  more 
productive  than  the  average  for  Kherson  oats  and  less  productive  for 
Red  Rustproof,  as  is  indicated  by  the  fact  that  each  variety  of  check 
was  considerably  more  effective  in  the  adjustment  of  the  yields  of 
strains  of  the  same  variety  than  of  strains  of  the  other.  But  neither 
check  was  a very  accurate  measure  of  the  productivity  of  the  soil, 
even  for  its  own  variety,  as  indicated  by  the  failure  of  adjustment  to 
reduce  variability  consistently  even  when  Kherson  strains  were  ad- 


Experiments  in  Field  Plot  Technic 


71 


justed  according  to  the  Kherson  check  and  Red  Rustproof  strains 
according  to  the  Red  Rustproof  check. 

Value  and  Limitations  of  Adjusting  Yields  by  Means  of  Check 
Plots. — The  effect  on  plots  variability  of  adjusting  yields  by  means  of 
check  plots  in  all  of  the  tests  is  shown  in  summary  form  in  Table  40. 
The  variability  of  the  test  plots  was  reduced  by  adjustment  in  three 
tests  and  was  increased  in  the  other  five.  It  is  noteworthy  that  the 
three  tests  in  which  plot  variability  was  reduced  by  adjustment  were 
characterized  by  high  plot  variability,  as  indicated  by  the  standard 
deviation  of  check  plots,  while  the  tests  in  which  adjustment  was  not 


Table  40. — Summary  of  Relative  Variability  of  Actual  and  Adjusted 
Yields  of  Interior  Rows  in  All  Tests. 


Test 

Season 

Number 
of  var- 
eties  or 
strains 

Number 
or  rep- 
lica- 
tions 

Average 

Actual 

yields 

% 

deviation 

Adjusted 

yields 

% 

Barley  Variety 

1919 

27 

3 

15.35 

12.91 

Oats  Variety 

1919 

24 

3 

10.23 

8.27 

Oats  Strain 

1919 

15 

4 

5.96 

6.80 

Wheat  Variety 

1920 

94 

4 

24.27 

15.32 

Wheat  Variety 

1921 

94 

Q 

o 

10.45 

11.52 

Wheat  Mixture 

1921 

30 

4 

9.84 

13.81 

Oats  Variety 

1921 

32 

4 

10.86 

11.43* 

Oats  Strain 

1921 

64 

4 

10.82 

11.07* 

* Adjustment  by  Kherson  check. 


effective  were  in  general  low  in  plot  variability.  In  1919  adjustment 
was  quite  effective  in  reducing  variability  in  the  oats  variety  test,  while 
it  increased  variability  in  the  oats  strain  test,  which  was  conducted  on 
the  same  field  and  similarly  handled  in  every  way.  In  fact,  conditions 
were  considered  more  favorable  for  the  effectiveness  of  the  practice 
in  the  strain  test  than  in  the  variety  test,  for  the  check  plots  were  closer 
together  and  12  of  the  15  strains  tested  were  taxonomically  identical 
with  the  check.  But  the  standard  deviation  of  check  plots  on  the 
part  of  the  field  on  which  varieties  were  grown  was  almost  twice  as 
great  as  on  the  part  of  the  field  on  which  the  strains  were  grown.  Ap- 
parently the  high  variability  of  the  plots  in  the  variety  test  was  caused 
in  large  part  by  differences  in  actual  soil  productivity  which  were 
largely  counteracted  by  the  adjustment  of  yields,  while  there  was 
little  variation  in  soil  productivity  in  the  strain  test  and  such  plot 
variability  as  occurred  was  largely  due  to  other  factors.  In  general 
therefore  the  adjustment  of  yields  will  probably  be  found  more  ef- 


72 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


fective  on  fields  highly  variable  in  soil  productivity  than  on  more 
uniform  fields,  and  for  similar  reasons  the  method  will  probably  be 
found  more  effective  in  tests  covering  a rather  large  area  than  in  tests 
covering  a smaller  area. 

It  is  clear  that  the  adjustment  of  yields  by  means  of  check  plots 
entails  several  serious  disadvantages,  and  may  increase  experimental 
error  considerably.  Not  only  is  the  yield  of  the  check  plot  a far  from 
perfect  measure  of  soil  productivity  for  the  check  variety,  but  the  pro- 
ductivity of  the  same  soil  for  other  varieties  may  be  decidedly  dif- 
ferent. The  method  is  therefore  more  effective  in  tests  of  strains  of 
the  same  variety  as  the  check,  than  in  tests  of  different  varieties.  When 
the  yields  of  check  plots  are  materially  affected  by  factors  not  similarly 
affecting  the  neighboring  test  plots,  adjustment  of  yields  will  increase 
experimental  error.  The  check  plots  must  therefore  be  effectively 
protected  from  competition,  border  effect,  mechanical  errors,  and  the 
like.  Moreover,  it  is  to  be  expected  that  the  effectiveness  of  adjusting 
yields  will  vary  with  the  season,  since  the  relative  influence  of  soil 
productivity  on  yield  varies  with  the  season.  For  example,  in  a season 
in  which  winter  injury  is  exceptionally  severe,  actual  soil  fertility  may 
have  comparatively  little  to  do  with  plot  yields.  Now,  if  the  check 
variety  is  hardy,  its  yields  may  vary  with  the  soil  fertility,  but  when 
corresponding  adjustments  are  made  on  the  yields  of  tested  varieties 
limited  in  yield  by  winter  injury,  a decrease  in  the  variability  of  repli- 
cate plots  is  hardly  to  be  expected.  The  same  considerations  apply 
of  course  to  yields  limited  by  many  other  factors. 

But,  although  a multitude  of  objections  may  be  made  to  the 
theoretical  bases  of  the  practice  of  adjusting  yields  in  variety  tests, 
and  although  in  many  cases  it  undoubtedly  results  in  an  increase  rather 
than  a decrease  in  experimental  error,  the  practice  offers  promise  of 
value  and  is  worthy  of  further  investigation.  The  effectiveness  of 
the  adjustment  of  yields  in  the  wheat  variety  test  of  1920,  in  which 
the  variability  of  replicate  test  plots  was  reduced  about  40  per  cent, 
is  a demonstration  of  the  possibilities  of  the  method.  An  increase  in 
replication  of  plots  involving  the  same  increase  in  land  and  labor  would 
probably  have  reduced  plot  variability  only  about  7 per  cent.  A thor- 
ough knowledge  of  the  value  and  limitations  of  yield-adjustment  by 
means  of  check  plots  might  enable  us  to  reduce  variability,  at  least  in 
some  types  of  plot  tests,  much  more  effectively  by  this  means  than 
by  replication.  The  saving  in  area  required  is  of  particular  significance 
in  preliminary  tests  if  border  rows  must  be  used  for  the  elimination 
of  competition,  since  in  this  case  the  area  required  for  a large  number 
of  replications  is  in  many  cases  prohibitive. 


Experiments  in  Fieed  Plot  Technic 


73 


CONCLUDING  REMARKS 

The  best  method  for  preliminary  variety  testing  is  one  which  will 
permit  the  accurate  determination  of  the  relative  value  of  the  va- 
rieties under  field  conditions,  with  the  use  of  a small  area  of  land  for 
each  variety.  Some  precision  must  be  sacrificed  to  save  land,  and  in 
so  far  as  the  errors  involved  are  of  such  nature  that  their  extent  can 
be  approximately  determined,  and  conclusions  drawn  accordingly,  this 
sacrifice  of  precision  is  permissible.  In  many  cases  it  is  advisable,  for 
example,  to  reduce  the  number  of  replications  and  to  increase  the  least 
difference  in  yield  regarded  significant  to  a sufficient  degree  to  com- 
pensate for  the  decrease  in  precision. 

But  these  considerations  do  not  apply  to  systematic  errors,  which, 
since  they  affect  the  yields  of  replicate  plots  similarly,  and  consequently 
have  little  effect  on  plot  variability,  cannot  be  accurately  measured. 
Typical  systematic  errors  commonly  involved  in  preliminary  testing 
are  (1)  modification  of  growing  conditions  favoring  some  varieties 
more  than  others,  such  as  hand  planting  or  wide  spacing  between  rows, 
and  (2)  competition  between  varieties  of  different  type,  resulting  from 
the  use  of  single-row  plots.  The  relative  value  of  varieties  under 
such  conditions  may  be  vastly  different  from  their  relative  value  un- 
der typical  field  conditions.  Even  should  measurable  experimental 
error  be  reduced  to  the  absolute  minimum,  such  a variety  test  might 
give  results  entirely  misleading.  The  error  cannot  be  counteracted, 
as  can  non-systematic  errors,  by  increasing  the  least  difference  con- 
sidered significant,  nor  can  the  extent  of  error  of  this  sort  be  measured 
or  estimated  by  a study  of  the  experimental  results. 

Systematic  error  must  therefore  be  reduced  by  every  practicable 
means.  Growing  conditions  in  the  preliminary  test- should  be  made  as 
similar  to  ordinary  field  conditions  as  possible.  The  effect  of  varietal 
competition  must  be  reduced  to  the  minimum.  If  this  can  be  ac- 
complished without  increasing  the  size  of  plots,  it  is  desirable  to  do 
so.  On  the  other  hand,  if  larger  plots  are  necessary  for  the  control 
of  competition,  larger  plots  should  be  used.  If  the  area  to  be  used 
for  preliminary  testing  cannot  be  correspondingly  increased,  the  num- 
ber of  replications  can  be  reduced  sufficiently  to  permit  the  use  of 
the  larger  plots  required  on  the  area  available.  This  will  necessitate 
a decrease  in  the  degree  of  precision  of  the  test,  and  will  reduce  the 
rapidity  of  elimination  of  the  less  valuable  varieties.  But  is  it  not 
better  to  eliminate  the  undesirable  varieties  slowly  than  to  risk  the 
elimination  of  desirable  ones  by  a more  rapid  analysis? 


74 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


The  error  from  competition  is  greater  when  different  varieties 
are  compared  than  when  different  strains  of  the  same  variety  are 
compared,  and  the  extent  of  error  is  roughly  in  proportion  to  the  de- 
gree of  difference  in  type  of  the  varieties  tested.  Competition  was 
not  found  to  be  correlated  closely  enough  with  earliness  of  heading, 
earliness  of  maturity,  height,  or  grain-straw  ratio  in  these  experi- 
ments to  permit  its  control  by  grouping  varieties  in  respect  to  these 
characters.  The  factor  found  most  closely  correlated  with  competitive 
value  was  yield,  but  the  correlation  even  in  this  case  was  not  close 
enough  to  permit  of  effective  control  by  grouping  varieties.  Moreover, 
it  would  be  impossible  in  practice  to  group  varieties  with  regard  to 
yield,  since  the  relative  yield  of  varieties  varies  so  widely  with  the 
season.  The  variety  expected  to  yield  poorly  is  not  ordinarily  included 
in  the  variety  test. 

When  different  strains  of  the  same  variety  are  grown,  the  error 
from  competition,  in  some  cases  at  least,  may  be  slight  enough  to 
justify  the  use  of  single-row  plots.  However,  competition  in  such 
cases  is  not  wholly  absent,  and  may  occasionally  be  quite  marked.  The 
importance  of  competition  as  a source  of  error  in  tests  of  pure  line 
selections  of  the  same  variety  merits  detailed  investigation.  If  it  is 
found  that  the  effects  of  competition  between  pure  lines  is  slight  it 
may  be  practicable  to  use  single-row  plots,  or  at  any  rate  to  use  3-row 
plots  without  discarding  border  rows.  The  latter  method  will  reduce 
the  error  from  competition  materially,  without  necessitating  the  loss 
of  any  of  the  experimental  area.  When  the  same  total  area  is  used, 
however,  single  row  plots  are  somewhat  more  reliable  than  3-row 
plots,  because  more  replications  can  be  used.  The  best  size  of  plot  for 
ordinary  variety  testing,  as  indicated  by  this  investigation,  is  probably 
the  3-row  plot  with  border  rows  discarded.  The  length  of  the  plot 
as  harvested  is  assumed  to  be  16  feet,  but  the  same  considerations  will 
apply  for  any  other  convenient  length.  The  number  of  replications 
will  vary  with  the  heterogeneity  of  the  field  and  the  degree  of  precision 
required  (and,  to  some  extent,  with  the  season  and  the  variety). 

Check  plots  have  been  used  in  preliminary  variety  tests  mainly 
for  the  following  purposes: 

(1)  For  the  adjustment  of  the  yields  of  the  test  plots,  and 

(2)  To  provide  a measure  of  plot  variability  for  the  field  used, 
and  thus  to  determine  the  degree  of  precision  of  the  experimental  re- 
sults, or  the  number  of  replications  which  would  be  required  for  a 
given  degree  of  precision. 

In  both  cases  the  behavior  of  the  check  variety  is  the  basis  for 
conclusions  regarding  the  tested  varieties.  This  involves  the  as- 
sumption that  different  varieties  of  the  same  crop  respond  similarly 


Experiments  in  Field  Plot  Technic 


75 


to  varying  conditions.  In  one  case,  reported  in  this  paper,  two  stand- 
ard varieties,  used  as  duplicate  checks,  and  grown  side  by  side  in 
120  distributed  sections  of  a field,  showed  no  significant  correlation 
in  relative  yield  of  adjoining  plots,  and  differed  so  widely  in  plot  varia- 
bility that  the  number  of  replications  necessary  for  a given  degree  of 
accuracy  was  more  than  twice  as  great  for  one  check  variety  as  for  the 
other.  Further  investigation  is  necessary  to  determine  how  generally 
such  cases  may  occur,  but  this  single  case  indicates  at  least  a possible 
source  of  extreme  error  in  the  use  of  check  plots,  either  for  adjust- 
ment of  yield  or  for  the  determination  of  the  probable  error  of  the 
experimental  results. 

For  this  and  various  other  reasons  the  adjustment  of  yields  by 
means  of  check  plots  is  at  present  of  doubtful  value  as  a general  prac- 
tice. In  some  cases,  however,  such  adjustment  accomplishes  a great 
improvement  in  the  precision  of  an  experiment,  with  a relatively  slight 
increase  in  expense.  The  practice  is  more  promising  for  tests  of 
strains  or  selections  of  the  same  variety  than  for  tests  of  different 
types.  A thorough  study  of  the  use  of  check  plots  in  variety  and  strain 
testing  may  discover  methods  of  overcoming  the  disadvantages,  and 
thus  make  available  an  economical  and  effective  method  of  increasing 
precision.  Meanwhile,  check  plots  should  be  used  cautiously.  Meth- 
ods for  adjusting  yields  and  for  determining  the  extent  of  plot  varia- 
bility without  the  use  of  check  plots  are  available17’ 18,  and  check 
plots  must  demonstrate  actual  value  if  they  are  to  continue  in  use  in 
variety  tests. 


SUMMARY 

1.  In  variety  and  strain  tests  of  barley,  oats,  and  wheat,  in  five- 
row  blocks,  the  competing  border  rows  of  adjacent  sorts  gave  relative 
yields  often  widely  different  from  those  of  the  interior  rows  of  the 
same  plots. 

2.  Such  competitive  effects  were  much  more  extreme  between 
different  varieties  than  between  different  commercial  strains  of  the 
same  variety. 

3.  A considerable  error  from  competition  affected  tests  in  rows 
running  north  and  south,  as  well  as  those  in  rows  running  east  and 
west. 

4.  Although  in  general  the  higher  yielding  varieties  were  favored 
in  competition,  the  reverse  frequently  occurred.  In  some  cases  a ma- 
terial advantage  in  yield  in  the  interior  rows  was  converted  to  a 
material  disadvantage  in  yield  in  the  border  rows. 


76 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


5.  Competing  quality  was  correlated  fairly  consistently  with 
yield  and  with  earliness  of  heading  and  maturity.  No  relation  to 
grain-straw  ratio  was  found  in  the  one  season  in  which  this  charac- 
ter was  determined.  A significant  correlation  between  competition 
and  height  was  found  in  the  wheat  variety  test  of  1921,  but  the  rela- 
tion of  competition  to  height  was  not  determined  in  the  other  tests. 

6.  In  the  oats  tests  competition  was  most  closely  related  to  earli- 
ness of  heading  and  maturity,  but  was  also  related  to  yield.  In  the 
wheat,  competition  was  related  fairly  closely  to  both  yield  and  earliness. 
In  the  barley  it  was  not  significantly  correlated  with  any  of  the  char- 
acteristics studied,  though  the  relation  to  yield  was  considerably  closer 
than  the  relation  to  any  of  the  other  characteristics. 

7.  In  the  wheat  and  oats  tests  in  which  earliness  and  yield  were 
correlated  with  competition,  earliness  and  yield  were  correlated  quite 
closely  with  one  another. 

8.  Single-row  plots,  protected  from  competition  by  border  rows 
discarded  at  harvesting,  were  somewhat  more  variable  in  yield  than 
3-row  plots  similarly  protected,  but  the  difference  was  not  great 
enough  to  outweigh  their  advantage  in  size.  The  mean  yield  of  five 
replicate  protected  single-row  plots  is  therefore  more  reliable,  under 
the  conditions  of  these  tests,  than  the  mean  yield  of  three  replicate 
protected  3-row  plots,  which  would  occupy  the  same  area  and  require 
considerably  more  labor  in  harvesting  and  threshing. 

9.  There  was  no  consistent  difference  in  variability  between  3- 
row  and  5-row  plots. 

10.  Plot  variability  was  increased  with  increase  in  the  size  of  the 
experiment  field.  The  number  of  replications  required  for  a given 
degree  of  precision,  as  measured  by  the  variability  of  plot  yields,  is 
therefore  increased  somewhat  when  border  rows  are  added  for  the 
control  of  competition. 

11.  The  variability  of  120  distributed  check  plots  of  Kherson  oats 
differed  widely  from  that  of  120  distributed  plots  of  Red  Rustproof 
oats,  adjacent  to  them.  If  the  variability  of  the  check  yields  were  con- 
sidered a measure  of  the  precision  of  the  test,  entirely  different  con- 
clusions would  be  drawn  on  the  basis  of  the  yields  of  these  two  check 
varieties. 

12.  Adjustment  of  plot  yields  on  the  basis  of  the  yields  of  check 
plots  resulted  in  a decrease  in  plot  variability  in  three  tests  and  in  an 
increase  in  five  tests.  In  general  the  practice  was  effective  on  fields  of 
high  plot  variability,  and  was  ineffective  on  fields  of  low  plot  varia- 
bility. 


Experiments  in  Field  Plot  Technic 


77 


13.  In  the  oats  strain  test  in  which  both  Kherson  and  Red  Rust- 
proof check  plots  were  included,  the  Kherson  check  was  more  effect- 
ive than  the  Red  Rustproof  check  as  a basis  for  adjusting  the  yields  of 
the  Kherson  strains,  while  the  Red  Rustproof  check  was  more  ef- 
fective as  a basis  for  adjusting  the  yields  of  the  Red  Rustproof  strains. 

14.  The  correlation  between  the  yields  of  adjacent  Kherson  and 
Red  Rustproof  check  plots  was  not  statistically  significant.  Adjust- 
ment of  the  yields  of  the  Kherson  check  plots  on  the  basis  of  the 
yields  of  the  adjacent  Red  Rustproof  plots,  and  of  those  of  the 
Red  Rustproof  plots  on  the  basis  of  the  Kherson  yields  increased  va- 
riability. 


ACKNOWLEDGMENT 

The  writer  is  indebted  to  Professors  M.  F.  Miller  and  W.  C. 
Etheridge  for  a critical  reading  of  the  manuscript,  and  to  O.  W. 
Letson  for  preparing  figure  8. 


78 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  49 


REFERENCES  CITED. 

1.  Day,  James  W.  The  relation  of  size,  shape,  and  number  of  replications 

of  plats  to  probable  error  in  field  experimentation.  In  Journ.  Amer.  Soc. 
Agron.  12,  3;  pp.  100-105.  1920. 

2.  Etheridge,  W.  C.  A classification  of  the  varieties  of  cultivated  oats.  Cor- 

nell Univ.  Agr.  Expt.  Sta.  Memoir  10 ; pp.  85-172.  1916. 

3.  Hall,  A.  D.  and  E.  J.  Russell.  Field  trials  and  their  interpretation.  In 

Jour.  Bd.  Agr.  (London)  Supplement:  pp.  5-14.  1911. 

4.  Hayes,  H.  K.  and  A.  C.  Arny.  Experiments  in  field  technic  in  rod-row 

tests.  In  Jour.  Agr.  Res.,  11,  9:  pp.  399-419.  1917. 

5.  Kiesselbach,  T.  A.  Studies  concerning  the  elimination  of  experimental  error 

in  comparative  crop  tests.  Nebr.  Agr.  Expt.  Sta.  Res.  Bui.  13:  pp.  3-95. 
1918. 

6.  Kiesselbach,  T.  A.  Experimental  error  in  field  trials.  In  Journ.  Amer.  Soc. 

Agron.  11,  6:  pp.  235-241.  1919. 

7.  Kiesselbach,  T.  A.  Plat  competition  as  a source  of  error  in  crop  tests.  In 

Journ.  Amer.  Soc.  Agron.  11,  6 : pp.  242-247.  1919. 

8.  Love,  H.  H.  The  experimental  error  in  field  trials.  In  Journ.  Amer.  Soc. 

Agron.  11,  5:  pp.  212-216.  1919. 

9.  Love,  H.  H.  and  W.  T.  Craig.  Methods  used  and  results  obtained  in  cereal 

investigations  at  the  Cornell  Station.  In  Journ.  Amer.  Soc.  Agron.  io, 
4:  pp.  145-157.  1918. 

10.  Lyon,  T.  L.  A comparison  of  the  error  in  yield  of  wheat  from  plats  and 

from  single  rows  in  multiple  series.  In  Proc.  Amer.  Soc.  Agron.  2:  pp. 
38,  39.  1911. 

11.  Lyon,  T.  L.  Some  experiments  to  estimate  errors  in  field  plat  tests.  In 

Proc.  Amer.  Soc.  Agron.  3:  pp.  89-114.  1912. 

12.  Mercer,  W.  B.  and  A.  D.  Hall.  The  experimental  error  in  field  trials.  In 

Journ.  Agr.  Sci.  4,  2 : pp.  107-132.  1911. 

13.  Montgomery,  E.  G.  Variation  in  yield  and  methods  of  arranging  plats  to 

secure  comparative  results.  In  25th  Ann.  Rpt.  Nebr.  Agr.  Expt.  Sta. : 
pp.  164-180.  1911. 

14.  Montgomery,  E.  G.  Experiments  in  wheat  breeding.  Experimental  error 

in  the  nursery  and  variation  in  nitrogen  and  yield.  U.  S.  Dept.  Agr.  Bur. 
Plant  Indus.  Bui.  269 : pp.  5-61.  1913. 

15.  Morgan,  J.  O.  Some  experiments  to  determine  the  uniformity  of  certain 

plats  for  field  tests.  In  Proc.  Amer.  Soc.  Agron.  1 : pp.  58-67.  1910. 

16.  Salmon,  C.  Check  plats  as  a source  of  error  in  varietal  tests.  In  Journ. 

Amer.  Soc.  Agron.  6,  3:  pp.  128-131.  1914. 

17.  Surface,  F.  M.  and  Raymond  Pearl.  A method  for  correcting  for  soil 

heterogeneity  in  variety  tests.  In  U.  S.  Dept.  Agr.  Journ.  Agr.  Res.  5, 
22:  pp.  1039-1049.  1916. 

18.  Wood,  T.  B.  & F.  J.  M.  Stratton.  The  interpretation  of  experimental 

results.  In  Journ.  Agr.  Sci.  3,  4:  pp.  417-440.  1910. 


UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 

AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  50 


Certain  Responses  of  Apple  Trees  to 
Nitrogen  Applications  of  Different 
Kinds  and  at  Different  Seasons 

(Publication  Authorized  December  8,  1921) 


COLUMBIA,  MISSOURI 
JANUARY,  1922 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 

E.  EANSING  RAY,  P.  E.  BURTON,  H.  J.  BLANTON, 

St.  Eouis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 

J.  C.  JONES,  PH.  D.,  LL.  D.,  PRESIDENT  OF  THE  UNIVERSITY 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 

STATION  STAFF 

January,  1922 


AGRICULTURAL  CHEMISTRY 

C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  A.  M. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  SiEveking,  B.  S.  in  Agr. 

E.  M.  Cowan,  A.  M. 

AGRICULTURAL  ENGINEERING 

J.  C.  Wooley,  B.  S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr, 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

E.  F.  Hopkins,  Ph.  D. 

DAIRY  HUSBANDRY 

A.  C.  Ragsdale,  B.  S.  in  Agr. 

W.  W.  Swett,  A.  M. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B^S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  R.  McBride,  B.  S.  in  Agr. 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  A.  M. 

O.  W.  Letson,  A.  M. 

B.  M.  King,  B.  S.  in  Agr. 

Alva  C.  Hill,  B.  S.  in  Agr. 

Miss  Bertha  Hite,  A.  B.*  Seed  Analyst 
Miss  Pearl  Drummond,  A.  A.* 


RURAL  LIFE 
O.  R.  Johnson,  A.  M. 

S.  D.  Gromer,  A.  M. 

E.  L.  Morgan,  A.  M. 

B.  H.  Frame,  B.  S.  in  Agr. 


HORTICULTURE 

V.  R.  Gardner,  M.  S.  A. 

H.  D.  Hooker,  Jr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  M.  S. 

F.  C.  Bradford,  M.  S. 

H.  G.  Swartwout,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY 
H.  L-  Kempster,  B.  S. 

Earl  W.  Henderson,  B.  S. 

SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M. 

W.  A.  Albrecht,  Ph.  D. 

F.  L.  Duley,  A.  M.t 

R.  R.  Hudelson,  A.  M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr. 
Richard  Bradfield,  A.  B. 

O.  B.  Price,  B.  S.  in  Agr. 

VETERINARY  SCIENCE 
J.  W.  Connaway,  D.  V.  S.,  M.  D. 
L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 


OTHER  OFFICERS 
R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Secretary 
Sam  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 
Miss  Jane  Frodsham,  Librarian 

E.  E.  Brown,  Business  Manager 


*In  the  service  of  U.  S.  Department  of  Agriculture. 
tOn  leave  of  absence. 


Certain  Responses  of  Apple  Trees  to  Nitrogen 
Applications  of  Different  Kinds  and  at 
Different  Seasons. 

H.  D.  Hooker,  Jr. 

A careful  study  of  the  literature  dealing  with  fertilizer  appli- 
cations to  fruit  plants  shows  that  those  most  commonly  effective 
involve  nitrogen.  Other  elements  have  been  shown  to  be  beneficial 
in  many  cases  when  applied  to  fruit  plants,  but  not  so  consistently 
nor  so  strikingly.  The  relatively  distinct  effects  that  follow  occa- 
sional applications  of  iron  constitute  an  exception  to  the  general 
statement,  but  only  because  the  symptoms  of  iron  deficiency  are 
easily  recognized  and  generally  understood.  For  this  reason,  iron 
is  seldom  used  as  a fertilizer  except  where  its  requirement  is  clearly 
indicated  by  chlorosis.  Were  it  possible  to  tell  as  readily  when  the 
other  essential  mineral  elements  become  limiting  factors  of  growth 
and  production,  the  problem  of  fertilizer  requirements  would  be 
relatively  simple.  In  the  absence  of  any  symptoms  more  marked 
than  a pale  green  color  of  the  leaves  or  a small  amount  of  new 
growth — conditions  that  may  result  from  any  one  of  a number  of 
different  causes — it  is  inevitable  that  many  fertilizer  applications 
should  be  without  important  effect,  for  they  meet  no  requirement. 
On  the  whole,  it  is  remarkable  that  nitrogen  applications  should 
be  effective  in  increasing  growth  and  crop  producing  power  as 
generally  as  they  do,  a fact  which  indicates  that  nitrogen  is  more 
often  the  limiting  factor  of  growth  and  yield  in  fruit  trees  than 
any  of  the  essential  mineral  elements. 

Nitrogenous  fertilizers  applied  to  fruit  trees  have  quite  general- 
ly increased  the  set  of  fruit,  favored  vegetative  growth  and  increas- 
ed yields.  The  experimental  work  by  which  these  facts  have  been  de- 
termined has  been  for  the  most  part  empirical.  The  orchard  ferti- 
lizer problem  has  been  attacked  as  a simple  matter  of  generally  in- 
creasing growth  and  productiveness.  These  are,  to  be  sure,  the 
objects  of  ultimate  interest  and  of  greatest  importance,  but  the 
fertilizer  problem  is  more  complex.  The  response  in  terms  of 
growth  and  yields  is  the  culmination  of  many  different  activities — 
absorption,  elaboration,  utilization  and  storage — of  correlative  ef- 
fects on  other  constituents,  of  distinct  processes  of  growth,  fruit 
bud  differentiation,  fruit  setting  and  development,  each  of  which  is 
conditioned  by  different  factors  or  sets  of  factors.  For  example: 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  50 


Have  the  increased  yields  following  the  application  of  nitrogenous 
fertilizers  resulted  only  from  the  better  set  of  fruit  and  the  in- 
creased size  of  the  tree,  or  have  nitrogen  applications  had  a direct 
or  indirect  influence  on  fruit  bud  differentiation?  Little  attention 
has  been  given  the  effects  of  fertilizer  applications  made  at  times 
other  than  early  spring.  Is  the  season  of  application  which  is  best 
for  increasing  the  set  of  fruit  likewise  best  for  increasing  fruit  bud 
differentiation?  The  possible  advantages  to  be  derived  from  ferti- 
lizer applications  evidently  have  not  been  exhausted. 

The  fertilizer  problem  should  be  studied  as  a problem  in  nu- 
trition and  the  effects  of  nitrogen  applications  should  be  measured 
in  terms  of  the  chemical  changes  produced  in  various  parts  of  the 
plant  as  well  as  in  terms  of  growth  and  yield.  Only  by  this  method 
can  principles  of  more  or  less  general  significance  be  determined. 
The  following  questions  present  themselves:  Is  the  nitrogen  con- 
tent of  a fruit  tree  increased  by  nitrogen  applications?  Does  one 
form  of  nitrogenous  fertilizer  produce  a greater  immediate  effect 
than  another?  Is  the  content  of  other  constituents,  particularly  var- 
ious forms  of  carbohydrate,  altered?  Are  significant  changes  evident 
at  the  time  of  fruit  bud  differentiation  and  are  different  effects 
produced  by  applications  at  different  seasons?  In  the  hope  of 
throwing  light  on  some  of  these  questions,  the  present  investiga- 
tion was  begun  in  the  spring  of  1920. 

MATERIALS  AND  METHODS 

Through  the  kindness  of  Dr.  J.  C.  Jones,  President  of  the 
University  of  Missouri,  two  apple  orchards  at  McBaine,  Mis- 
souri, were  placed  at  the  disposal  of  the  Department  of  Horticul- 
ture for  experimental  treatment  and  sampling.  Each  orchard  con- 
tained York  trees  in  good  condition ; those  in  the  south  orchard 
were  20  years  old  and  bore  in  even  years;  those  in  the  north  or- 
chard were  16  years  old  and  bore  in  odd  years.  These  trees  were 
fertilized  in  a manner  to  be  described  presently  and  samples  of  spurs 
and  of  bark  from  the  scaffold  limbs,  four  to  six  inches  in  diameter, 
were  collected  at  intervals  during  the  year.  The  bark  samples  con- 
sisted of  strips  three-quarters  of  an  inch  wide  and  six  to  eight 
inches  long,  pointed  at  each  end  and  including  all  tissue  outside  of 
the  cambium.  Not  more  than  three  strips  were  taken  from  a sin- 
gle tree  at  one  sampling.  The  wounds  were  covered  with  white 
lead  paint  to  prevent  infection  and  undue  water  loss.  The  spur 
samples  included  1919  and  1920  growth. 


Responses  of  Apple  Trees  to  Nitrogen  Applications  5 

The  other  trees  used  for  experimental  purposes  were  at  the 
Experiment  Station  Fruit  Farm,  Turner  Station,  Mo.  Samples  of 
spurs  including  1919  and  1920  growth  were  collected  from  ferti- 
lized and  check  trees  of  Jonathan  and  Ben  Davis  varieties  growing 
in  bottom  land  and  seven  years  old  in  1920,  when  the  samples  were 
collected.  Another  fertilizer  experiment  was  conducted  on  vigor- 
ous four-year-old  Grimes  trees  at  the  Fruit  Farm.  Samples  of 
spurs  of  various  lengths  were  collected  from  mature  Ben  Davis 
trees  growing  in  the  University  orchard  at  Columbia. 

The  chemical  analyses  were  made  after  the  manner  detailed 
in  Research  Bulletin  40  of  this  Station.  Determinations  were  made 
of  dry  weight,  ash,  potassium,  phosphorus,  nitrogen,  reducing  sug- 
ars, total  sugars  and  starch. 

In  the  spring  of  1921  practically  all  blossoms  in  the  orchards 
under  study  were  killed  by  late  frosts.  Since  it  was  impossible  to 
determine  the  effects  of  all  the  fertilizer  treatments  as  originally 
planned,  some  of  the  treatments  were  repeated  in  1921.  However, 
a large  body  of  data  had  been  collected  which  it  seems  advisable 
to  publish,  incomplete  and  fragmentary  though  it  be,  since  it  shows 
some  significant  and  rather  unexpected  facts. 

SPRING  APPLICATIONS  OF  VARIOUS  NITROGENOUS 

FERTILIZERS 

This  first  experiment  was  made  on  York  trees  in  their  bearing 
year.  Four  plots  of  15  trees  each  were  selected ; one  was  left  as  a 
check;  another  received  5 pounds  of  sodium  nitrate  per  tree;  an- 
other received  3 pounds  of  ammonium  sulphate  per  tree  and  the 
other  5 pounds  of  a high  grade  of  dried  blood.  By  this  treatment 
each  fertilized  tree  received  approximately  the  same  amount  of 
nitrogen.  The  applications  were  made  March  19,  1920. 

The  greater  crop  produced  by  the  fertilized  plots  was  the  most 
striking  effect  produced.  The  yields  from  the  plots  were  as  fol- 
lows: 285  bushels  from  the  check  plot  or  19.7  bushels  per  tree; 

375  bushels  from  the  blood  plot  or  25.0  bushels  per  tree;  381  bush- 
els from  the  nitrate  plot  or  25.4  bushels  per  tree;  376  bushels  from 
the  ammonium  sulphate  plot  or  25.1  bushels  per  tree.  These  fig- 
ures show  that  the  three  types  of  fertilizers  used  produced  practi- 
cally the  same  effect  in  increasing  yield.  There  were  minor  vari- 
ations within  the  plots:  one  tree  on  the  blood  plot  produced  40 
bushels  of  apples ; one  tree  on  the  ammonium  sulphate  plot  bore  so 
large  a crop  that  the  tree  split  to  the  ground  under  the  weight  of 


6 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  50 


fruit;  the  yields  of  the  trees  on  the  nitrate  plot  were  more  nearly 
uniform.  No  definite  figures  are  available  but  the  apples  from  the 
nitrate  plot  seemed  to  be  slightly  larger  and  somewhat  less  highly 
colored  than  those  from  the  other  plots. 

The  data  in  Table  1 show  that  neither  the  bearing  nor  the  non- 
bearing spurs  of  the  fertilized  trees  formed  a larger  number  of 
leaves  during  the  current  season  than  did  corresponding  spurs  on 
the  unfertilized  trees.  The  following  spring  there  was  no  bloom 
on  any  of  the  plots.  The  chief  effects  of  the  fertilizer  treatments 
observed  were  a deeper  color  of  the  foliage  and  an  increased  set  of 


Table  1. — Average  Number  oe  Leaves  Per  Spur  on  Fertilized  and  Unferti- 
lized York  Trees. 


May  3 

May  15 

May  22 

Bearing  spurs 

Check  plot  _ . 

7.0 

9.1 

9.25 

Nitrate  plot  — 

4.8 

7.2 

7.34 

Amm.  sulphate  plot  _ 

6.8 

8.4 

8.6 

Blood  plot  

5.8 

7.4 

7.9 

Non-Bearing  spurs 

Check  plot 

6.6 

8.0 

8.2 

Nitrate  plot 

5.2 

7.2 

7.4 

Amm.  sulphate  plot  __ 

6.0 

7.4 

7.6 

Blood  plot  

5.7 

7.1 

7.4 

fruit:  23.7  percent  of  the  blossoming  spurs  on  the  check  trees 

bore  fruit  and  approximately  32  percent  of  the  blossoming  spurs 
on  the  fertilized  trees,  as  determined  by  actual  count. 

Samples  of  spurs  and  of  bark  were  collected  from  the  trees 
on  these  plots  at  five  dates;  May  22,  June  19,  September  6,  Novem- 
ber 20,  1920,  and  April  2,  1921.  The  analytical  determinations  on 
these  samples  are  given  in  Table  2.  (See  pages  8 and  9.) 

The  greater  set  of  fruit  on  the  fertilized  trees  is  probably  to 
be  associated  with  the  greater  nitrogen  content  of  their  spurs  on 
May  22,  as  suggested  by  Harvey  and  Murneek  (Ore.  Agr.  Expt. 
Sta.  Bui.  176).  At  this  time  the  nitrogen  content  of  the  spurs  is 
on  the  decline,  as  reference  to  Figure  10  in  Research  Bulletin  40 
of  this  Station  shows. 

Except  for  this  temporary  increase  in  nitrogen  content  there 
is  no  very  significant  difference  between  the  composition  of  the 
fertilized  and  unfertilized  trees.  The  absence  of  such  variations 


Responses  of  Apple  Trees  to  Nitrogen  Applications 


7 


is  striking.  There  has  been  no  marked  starch  accumulation  in  any 
of  the  spurs  by  June  19,  immediately  before  the  period  of  fruit  bud 
differentiation.  Moreover,  on  April  2 the  following  spring,  the  spurs 
of  the  fertilized  trees  contained  very  little  more  nitrogen  than  the 
check  spurs,  the  difference  found  being  well  within  the  limits  of 
experimental  error.  Similarly,  though  the  nitrogen  content  of  the 
bark  from  the  nitrated  trees  is  slightly  greater  than  that  of  the 
check  trees,  the  sulphate  and  blood  trees  have  less.  Consequently 
no  consistent  difference  is  evident. 

The  comparison  between  the  chemical  composition  of  spurs 
and  bark  from  the  same  trees  as  afforded  by  the  data  presented  is 
of  considerable  interest.  In  general,  the  variations  in  the  chemical 
composition  of  the  bark  follow  rather  closely  those  in  the  spurs. 
The  low  starch  content  of  the  former  in  May  and  June  is  particu- 
larly striking  as  it  indicates  that  the  factors  which  prevent  carbo- 
hydrate accumulation  in  the  bearing  spurs  likewise  affect  the  bark 
of  the  scaffold  limbs.  The  high  total  sugar  content  of  the  bark 
during  the  winter  and  especially  in  April  is  apparently  character- 
istic. The  bark  contains,  during  most  of  the  year,  about  half  the 
percentage  of  nitrogen  that  the  spurs  contain ; its  percentage  phos- 
phorus content  is  also,  for  the  most  part,  much  less  but  the  per- 
centage potassium  content  of  these  two  portions  of  the  tree  is  of 
the  same  order  and  during  part  of  the  year  the  bark  contains  an 
even  higher  percentage  than  the  spurs.  This  is  consistently 
true  in  the  June  and  September  analyses.  The  percentage  ash 
content  of  bark  is  usually  greater  than  that  of  the  spurs  though  the 
difference  is  very  nearly  wiped  out  in  the  spring. 

In  this  experiment  the  various  types  of  nitrogenous  fertilizers 
have  produced  essentially  the  same  effects  as  shown  both  by  the 
chemical  analyses  and  the  crop  yields.  This  effect  has  consisted 
principally  in  an  increased  set.  There  has  been  no  effect  in  in- 
creasing the  number  of  leaves  during  the  current  season  and  very 
little  effect,  if  any,  on  the  rate  of  growth.  There  has  been  no  effect 
on  fruit  bud  differentiation  nor  any  tendency  in  that  direction,  such 
as  might  be  evidenced  by  an  accumulation  of  starch  in  the  spurs 
during  the  period  of  fruit  bud  differentiation.  In  fact,  the  greater 
carbohydrate  utilization  following  the  increased  set  of  fruit  would 
tend  to  decrease  the  chances  for  starch  accumulation  in  the  ab- 
sence of  an  increased  leaf  area  per  spur. 

Nitrogenous  fertilizers  should,  therefore,  be  applied  to  bien- 
nially bearing  apple  trees  in  the  spring  of  their  crop  year  with  ex- 


8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  50 


treme  caution,  for  such  a practice  is  likely  to  lead  to  overproduc- 
tion with  its  attendant  evils. 


Table  2. — Analyses  oe  Spurs  and  Baric  From  Fertilized  and  Unfertilized 
York  Trees  in  Their  Bearing  Year. 

( Percentages  of  dry  weight) 


Dry 

weight 

Reduc’g 

sugars 

Total 

sugars 

Starch 

Ash 

K 

P 

N 

May  22,  1920 

Bearing  spurs 

Check  

40.2 

0.79 

1.95 

0.68 

9.73 

0.678 

0.170 

1.020 

Nitrate  

43.2 

1.00 

1.65 

1.15 

8.74 

0.626 

0.162 

1.047 

Sulphate  __  _ 

43.6 

0.66 

1.59 

0.00 

11.32 

0.649 

0.138 

1.164 

Blood  

41.2 

0.70 

2.79 

0.45 

7.69 

0.665 

0.174 

1.221 

Bark 

Check  

43.4 

1.48 

1.74 

0.11 

7.97 

0.621 

0.060 

0.578 

Nitrate 

43.1 

1.61 

1.89 

0.20 

12.79 

0.648 

0.064 

0.565 

Sulphate 

42.7 

1.69 

1.74 

0.00 

8.12 

0.585 

0.072 

0.605 

Blood  

42.3 

1.53 

2.10 

0.14 

8.32 

0.513 

0.075 

0.526 

June  19,  1920 

Bearing  spurs 

Check  _ 

44.4 

1.26 

1.65 

0.00 

6.44 

0.554 

0.140 

0.916 

Nitrate  

45.7 

1.26 

1.29 

0.00 

6.59 

0.362 

0.141 

0.912 

Sulphate  _ _ 

42.5 

0.77 

1.08 

0.36 

6.02 

0.639 

0.148 

1.024 

Blood  _ 

43.1 

0.86 

1.20 

0.00 

7.21 

0.558 

0.209 

1.166 

Bark 

Check  

47.9 

1.32 

1.62 

0.00 

9.91 

0.570 

0.145 

0.578 

Nitrate  

42.3 

1.71 

2.00 

0.38 

8.41 

0.583 

0.108 

0.546 

Sulphate 

42.6 

1.62 

1.89 

0.70 

8.11 

0.647 

0.086 

0.575 

Blood 

48.1 

1.73 

2.04 

0.54 

7.77 

0.522 

0.096 

0.551 

September  6,  1920 

Bearing  spurs 

Check  _ 

46.7 

1.21 

2.25 

2.88 

7.97 

0.408 

0.155 

1.030 

Nitrate 

45.6 

0.88 

1.38 

1.98 

7.29 

0.371 

0.164 

0.880 

Sulphate  _ _ 

45.3 

0.72 

1.14 

2.41 

7.29 

0.420 

0.211 

1.110 

Blood  

48.8 

0.72 

1.20 

1.98 

6.58 

0.414 

0.176 

1.010 

Bark 

Check  

44.9 

1.87 

2.70 

2.53 

10.76 

0.457 

0.110 

0.52 

Nitrate  

40.6 

1.72 

2.60 

2.27 

11.45 

0.411 

0.107 

0.44 

Sulphate  

43.6 

1.65 

2.55 

2.36 

11.08 

0.502 

0.108 

0.58 

Blood  

44.6 

1.45 

1.86 

2.66 

12.00 

0.459 

0.136 

0.54 

November  20,  1920 

Spurs 

Check  

48.4 

2.95 

3.15 

1.08 

9.05 

0.489 

0.222 

1.22 

Nitrate  

47.9 

1.98 

2.73 

1.19 

8.01 

0.457 

0.230 

1.14 

Sulphate  _ 

49.6 

1.94 

2.71 

1.28 

8.53 

0.473 

0.177 

1.09 

Blood 

48.5 

1.73 

2.16 

1.15 

8.69 

0.513 

0.241 

1.20 

Responses  of  Apple  Trees  to  Nitrogen  Applications 


9 


Table  2. — (Continued.) 


Dry 

weight 

Reduc’g 

sugars 

Total 

sugars 

Starch 

Ash 

K 

P 

N 

Bark 

Check  

44.1 

2.68 

3.78 

1.58 

12.35 

0.498 

0.098 

0.69 

Nitrate  __ 

45.3 

3.07 

4.32 

1.72 

11.48 

0.393 

0.102 

0.61 

Sulphate  — 

46.9 

2.81 

3.42 

1.58 

12.33 

0.385 

0.075 

0.64 

Blood  

44.1 

2.38 

3.27 

1.98 

11.43 

0.397 

0.106 

0.58 

April  2,  1921 

Spurs 

Check  

48.5 

2.44 

3.06 

1.53 

9.28 

0.498 

0.210 

0.86 

Nitrate  

46.9 

2.36 

3.37 

2.34 

9.71 

0.428 

0.212 

0.92 

Sulphate  

47.2 

2.07 

3.21 

1.35 

9.81 

0.441 

0.183 

0.89 

Blood  

44.7 

2.47 

3.45 

1.71 

10.11 

0.490 

0.202 

0.88 

Bark 

Check  

46.1 

5.57 

6.56 

2.55 

9.22 

0.405 

0.122 

0.58 

Nitrate  

46.7 

5.93 

6.36 

2.52 

10.63 

0.354 

0.104 

0.62 

Sulphate  _ _ 

46.4 

5.68 

5.94 

2.34 

10.19 

0.346 

0.147 

0.52 

Blood  

47.1 

5.79 

6.00 

2.43 

9.63 

0.430 

0.144 

0.56 

EFFECT  OF  SPRING  APPLICATIONS  OF  NITRATE  IN 
PROMOTING  GROWTH 

Spring  applications  of  nitrogenous  fertilizers  not  only  have  an 
effect  on  the  setting  of  fruit,  but,  as  is  well  known,  they  frequently 
increase  the  amount  of  growth.  This  effect  is  shown  by  an  experi- 
ment on  Ben  Davis  and  Jonathan  trees  at  the  University  Fruit 
Farm.  Two  trees  of  each  variety  were  treated  with  three  pounds 
of  sodium  nitrate  in  the  spring  of  1919  and  again  March  29,  1920. 
The  effects  of  this  treatment  have  been  revealed  in  the  greater  size 
of  the  fertilized  trees  as  compared  with  check  trees  of  the  same 
variety  in  the  same  rows.  These  trees  blossomed  for  the  first  time 
in  1921  but  the  entire  bloom  was  killed  by  spring  frost. 

Samples  of  spurs  were  collected  from  the  fertilized  trees  and 
the  checks  at  three  dates,  March  29,  May  22  and  June  19,  1920. 
The  analyses  of  these  spurs  are  given  in  Table  3. 

The  percentage  nitrogen  content  of  the  spurs  from  the  ferti- 
lized trees  was  less  on  March  29  than  that  of  the  check  spurs,  show- 
ing that  the  fertilizer  applied  the  year  before  did  not  increase  the 
nitrogen  content  of  the  spurs.  In  May,  the  fertilized  spurs  con- 
tained a higher  percentage  of  nitrogen  than  the  checks ; but  since 
their  percentage  nitrogen  content  continued  to  decline  during  the 
month  following,  while  that  of  the  checks  increased,  the  fertilized 


10 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  50 


Table  3. — Analyses  oe  Spurs  on  Fertilized  and  Unfertilized  Jonathan  and 

Ben  Davis  Trees. 


( Percentages  of  dry  weight ) 


Dry 

weight 

Reduc’g 

sugars 

Total 

sugars 

Starch 

Ash 

K 

P 

N 

March  29,  1920 

Jonathan 
Nitrated  

1.64 

1.92 

1.35 

5.315 

0.489 

0.193 

1.23 

Check  



1.71 

1.77 

1.08 

6.480 

0.558 

0.220 

1.35 

Ben  Davis 
Check  

1.98 

2.13 

0.47 

8.020 

0.414 

0.190 

1.30 

Nitrated  

— 

1.31 

2.25 

0.72 

7.925 

0.402 

0.200 

1.245 

May  22,  1920 

Jonathan 
Check  

42.9 

1.04 

1.88 

1.01 

4.76 

0.722 

0.131 

0.857 

Nitrated 

40.9 

1.22 

2.34 

0.00 

4.70 

0.423 

0.127 

0.927 

Ben  Davis 
Check  

44.1 

1.04 

1.65 

0.36 

4.59 

0.514 

0.100 

0.699 

Nitrated 

43.9 

1.01 

1.77 

0.90 

4.33 

0.480 

0.103 

0.822 

June  19,  1920 

Jonathan 
Check  

45.1 

1.08 

2.67 

2.16 

4.64 

0.457 

0.157 

0.897 

Nitrated  __  . _ 

47.1 

0.70 

1.86 

1.60 

4.51 

0.455 

0.144 

0.820 

Ben  Davis 
Check  

49.3 

1.08 

1.38 

1.42 

5.94 

0.325 

0.224 

1.128 

Nitrated  

47.6 

0.65 

1.35 

1.10 

4.60 

0.321 

0.134 

0.792 

spurs  again  contained  less  than  the  checks  on  June  19.  This  slight 
difference  in  itself  may  not  be  particularly  significant  at  this  time, 
immediately  preceding  the  period  of  fruit  bud  differentiation,  but 
when  considered  in  its  relation  with  other  conditions  with  which 
it  is  associated  and  for  which  it  may  possibly  be  responsible,  it  may 
assume  great  importance.  At  this  time  the  starch  content  in  the 
spurs  of  the  fertilized  trees  was  distinctly  less  than  in  the  check 
spurs.  This  indicates  clearly  that  the  conditions  for  fruit  bud  dif- 
ferentiation were  not  improved  and  in  fact  were  made  less  favor- 
able by  the  spring  application  of  nitrate  of  soda.  The  smaller  ac- 
cumulation of  starch  in  the  spurs  of  fertilized  trees  immediately 
before  the  period  of  fruit  bud  differentiation  is  probably  related  to, 
if  not  actually  caused  by,  the  more  vigorous  growth  of  the  ferti- 
lized trees. 

It  is  evident  that  the  minimum  nitrogen  content  occurred 
much  sooner  in  the  spurs  of  the  check  trees  than  in  those  of  the 


Responses  of  Apple  Trees  to  Nitrogen  Applications  11 


fertilized  trees  and  this  minimum  is  related  to  the  time  of  growth 
cessation,  for  an  accumulation  of  nitrogen  as  shown  by  an  in- 
creased percentage  does  not  usually  occur  in  spurs  until  growth  has 
ceased.  The  figures  in  Table  3 show  that  the  potasssium  and  total 
ash  content  of  the  fertilized  spurs  is  consistently  less  than  in  the 
check  spurs  and  for  the  most  part  this  is  true  also  of  the  phosphorus 
content.  These  conditions  may  be  interpreted  as  further  conse- 
quences of  the  more  vigorous  growth  of  the  fertilized  trees  for  it  is  a 
general  rule  that  the  greater  the  length  of  the  spur  growth,  the  low- 
er is  its  ash  content  at  the  close  of  the  growing  season.  This  is 
shown  by  the  data  in  Table  4 which  are  analyses  of  Ben  Davis  spurs 
collected  May  21  and  July  2,  1920.  The  samples  were  collected  ac- 
cording to  spur  lengths  as  shown  in  this  table.  It  will  be  seen  that 
the  ash,  phosphorus  and  nitrogen  content  is  less,  the  longer  the 
spur  growth  of  the  current  season. 

Table  4. — Analyses  oe  Ben  Davis  Spurs  According  to  Length 
(1919  and  1920  wood  included) 

( Percentages  of  dry  weight ) 


Spur  length  Dry  Ash  KPN 

in  centimeters  weight 


May  21,  1920 


0.5-  1.0  48.8 

1.1-  1.5  45.6 

1.6-  3.0.  41.2 

3.1- 10.0  36.9 

July  2,  1920 

0.5-  1.0  48.4 

1.1-  1.5  47.3 

1.6-  3.0  46.9 

3.1- 10.0  46.8 


6.15 

0.597 

0.179 

0.891 

5.28 

0.568 

0.177 

0.876 

4.91 

0.536 

0.137 

0.705 

4.49 

0.572 

0.133 

0.700 

6.44 

0.513 

0.142 

0.754 

5.53 

0.531 

0.134 

0.754 

5.04 

0.527 

0.138 

0.726 

4.18 

0.475 

0.118 

0.684 

THE  EFFECT  OF  SPRING  APPLICATIONS  ON  IMMA- 
TURE TREES 

On  March  29,  1920,  two  plots  of  young  Grimes  apples  were 
fertilized,  one  with  2 pounds  of  nitrate  of  soda  and  one  with  2 
pounds  of  dried  blood  to  the  tree.  Each  plot  contained  10  trees 
and  a similar  block  of  10  trees  was  left  as  a check.  About  50  short, 
spur-like  growths  and  13  leaders  on  the  trees  of  each  plot  were 
labeled.  Growth  measurements  were  made  on  these  spur-like 
growths  and  on  the  four  shoots  arising  from  the  terminal  portion 


12 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  50 


of  each  leader.  The  effects  of  the  fertilizer  treatments  on  the 
growth  of  these  trees  is  shown  in  Table  5.  It  is  evident  that  the  ni- 
trate of  soda  had  a greater  effect  on  the  shoot  growth  than  did  the 
dried  blood.  Practically  no  difference  is  evident  between  the  rates 
of  growth  from  the  short  spur-like  branches. 

Chemical  analyses  of  these  two  types  of  growth  are  shown  in 
Table  6.  The  absence  of  any  marked  differences  between  the  va- 
rious plots  is  quite  striking.  Differences  do  appear,  however,  in 
the  chemical  composition  of  the  terminal  growth  as  compared  with 
that  of  the  shorter  spur-like  branches.  In  the  former  there  is  no 
complete  disappearance  of  starch  in  June  and  the  nitrogen  content 


Table  5. — Average  Lengths  in  Centimeters  oe  Growths  From  Leaders  and 
Spur-Like  Branches  on  Four-Year-Oed  Grimes. 


May  5 

May  14 

May  21 

May  31 

June  14 

July  9 

Leader  growth 

Check  

2.24 

8.5 

12.2 

19.5 

28.0 

35.8 

Nitrate  

2.23 

9.8 

14.7 

22.5 

31.9 

42.6 

Blood 

2.18 

8.7 

12.5 

19.0 

27.5 

34.6 

Twig  growth 

Check  _ 

2.4 

7.2 

9.4 

13.0 

15.3 

16.1 

Nitrate  

2.2 

7.5 

10.6 

13.9 

15.6 

16.0 

Blood  _ 2.7  9.3  10.4 

Average  number  oe  leaves  on  twig  growth 

13.3 

15.8 

16.8 

Check  _ _ 

6.0 

9.3 

10.0 

12.0 

13.9 

14.2 

Nitrate 

7.2 

8.9 

10.1 

11.8 

12.9 

13.1 

Blood 

7.5 

9.8 

10.8 

12.3 

13.5 

14.6 

rises  to  a high  maximum  in  May,  much  as  in  bearing  spurs  on  ma- 
ture trees. 

The  only  apparent  difference  associated  with  the  greater  ter- 
minal shoot  growth  of  the  nitrated  trees  is  shown  in  a higher  per- 
centage of  starch  and  total  sugar  and  a lower  percentage  of  ash, 
potassium  and  phosphorus  in  June.  In  September  these  shoots 
have  the  highest  percentage  of  ash  and  potassium  and  the  lowest 
of  phosphorus  and  nitrogen. 

The  data  presented  indicate  a differential  effect  between  ni- 
trate of  soda  and  dried  blood,  which  may  be  associated  with  the 
more  quickly  available  character  of  the  former.  Different  parts  of 
a tree  evidently  may  react  in  different  ways  to  the  same  fertilizer 
treatment.  A comparison  of  the  responses  of  these  four-year-old 
trees  with  those  of  the  Ben  Davis,  Jonathan  and  York  trees  indi- 


Responses  of  Apple  Trees  to  Nitrogen  Applications  13 


Table  6. — Analyses  oe  Growths  From  Leaders  and  Spur-Like  Branches  on 

Four- Year-Old  Grimes. 


( Percentages  of  dry  weight ) 


Dry 

weight 

Reduc’g 

sugars 

Total 

sugars 

Starch 

Ash 

K 

P 

N 

May  21,  1920 

Leader  growth 

Check  

25.8 

0.97 

1.17 

2.09 

4.49 

1.435 

0.234 

1.625 

Nitrate  

25.1 

0.77 

1.35 

2.05 

4.61 

0.669 

0.236 

1.625 

Blood  

25.2 

1.01 

1.26 

2.70 

4.56 

1.011 

0.216 

1.615 

Spur-like  growth 

Check 

31.0 

0.86 

1.44 

1.96 

4.63 

0.599 

0.204 

1.299 

Nitrate  

33.6 

0.90 

1.08 

1.51 

4.17 

0.706 

0.152 

1.045 

Blood  

33.0 

1.08 

1.08 

1.58 

4.26 

0.579 

0.181 

1.150 

June  19,  1920 

Leader  growth 
Check  

38.7 

0.88 

1.62 

2.18 

3.64 

0.791 

0.147 

0.845 

Nitrate 

38.5 

1.10 

1.95 

2.75 

3.08 

0.642 

0.137 

0.952 

Blood  

38.7 

1.28 

1.62 

2.12 

3.54 

0.697 

0.142 

1.063 

Spur-like  growth 

Check  

46.8 

0.95 

1.41 

0.02 

4.20 

0.578 

0.163 

0.739 

Nitrate  _ 

44.0 

0.90 

1.47 

0.00 

4.19 

0.689 

0.183 

0.829 

Blood  _ 

46.8 

1.13 

1.72 

0.00 

4.00 

0.299 

0.145 

0.675 

September  4,  1920 

Leader  growth 

Check  _ _ __ 

39.0 

0.54 

0.90 

1.40 

3.08  | 

; 0.319 

0.177 

0.76 

Nitrate 

39.6 

0.67 

1.05 

1.44 

3.55  1 

0.685 

0.150 

0.64 

Blood 

39.4 

0.29 

0.60 

1.73 

3.23  ! 

0.554 

0.153 

0.66 

Spur-like  growth 
Check  

53.0 

0.50 

0.75 

2.30 

5.3i 

0.433 

0.189 

0.75 

Nitrate  

53.0 

0.81 

1.23 

1.89 

4.98 

0.411 

0.197 

0.73 

Blood  

54.3 

0.68 

1.23 

2.05 

6.01 

0.406 

0.196 

0.81 

cates  that  the  age  of  the  tree  may  be  an  important  factor,  although 
it  is  quite  possible  that  the  observed  differences  may  have  been  due 
to  the  conditions  under  which  the  various  trees  were  growing. 

THE  EFFECT  OF  APPLICATIONS  AT  DIFFERENT 

SEASONS 

Fertilizer  applications  were  made  on  16-year-old  York  trees 
in  their  off  year.  One  plot  of  15  trees  was  fertilized  March  29  with 
dried  blood  and  another  on  June  20.  Both  times  5 pounds  of  fer- 
tilizer were  applied  to  each  tree.  On  September  20  another  plot 
of  15  trees  was  given  5 pounds  of  sodium  nitrate  to  the  tree.  Be- 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  50 


cause  of  the  severe  spring  frosts  of  1921,  the  effects  on  these  trees 
could  not  be  measured  except  in  terms  of  the  chemical  changes  ob- 
served in  the  bark  and  spurs.  These  are  given  in  Table  7. 

The  data  afford  a comparison  with  Table  1 of  the  chemical 
composition  of  bark  on  bearing  and  non-bearing  trees.  In  gener- 
al the  seasonal  changes  in  the  chemical  composition  of  the  bark 
on  the  scaffold  limbs  of  these  alternate  bearing  trees  follow  those 
of  the  spurs.  In  June  there  is  already  an  accumulation  of  carbo- 
hydrate in  the.  form  of  starch.  In  May  the  nitrogen  content  is 
somewhat  less  in  the  non-bearing  trees.  During  the  winter  the 
sugar  content  of  the  bark  is  exceptionally  high,  much  higher  in 
the  non-bearing  than  in  the  bearing  trees.  From  September 
through  March  the  potassium  content  of  the  non-bearing  trees  is 
distinctly  higher  and  the  phosphorus  and  nitrogen  content  like- 
wise, though  to  a lesser  extent. 

The  differential  effects  from  applying  these  fertilizers  at  va- 
rious seasons  is  most  evident  in  the  nitrogen  content  of  the  spurs. 
On  March  30,  1921,  the  nitrogen  content  of  the  spurs  varies  with 
the  lateness  of  application  the  previous  season.  The  check  spurs 
have  the  least,  the  spring  fertilized  trees  next,  the  summer  ferti- 
lized trees  more  and  the  fall  fertilized  trees  most  of  all.  Moreover 
these  differences  in  nitrogen  content  are  by  no  means  insignificant. 
It  is  unfortunate  that  weather  conditions  made  it  impossible  to 
follow  the  later  effects  associated  with  these  differences  in  the  ni- 
trogen content  of  the  spurs.  It  is  impossible  to  say  what  advan- 
tage might  accrue  from  increasing  the  nitrogen  content  of  spurs 
in  the  spring  of  their  bearing  year  but  it  is  clear  that  late  sum- 
mer or  early  fall  applications  of  nitrogenous  fertilizers  are  much 
more  effective  in  this  respect  than  spring  applications. 

The  effect  of  spring  application  of  blood  on  these  non-bearing 
trees  is  essentially  the  same  as  that  observed  on  the  Ben  Davis 
and  Jonathan  trees.  The  accumulation  of  starch  in  the  spurs  to- 
ward the  end  of  June  at  the  critical  time  of  fruit  bud  differentia- 
tion is  reduced  by  the  spring  application  of  dried  blood  just  as  it 
was  by  the  nitrate  of  soda  in  the  Ben  Davis  and  Jonathan  spurs. 
Moreover,  the  residual  effect  of  the  spring  fertilizer,  as  shown  by 
the  analyses  of  March  30,  1921,  is  practically  nil,  as  in  the  case  of 
the  bearing  York  trees. 

The  most  marked  effect  on  the  nitrogen  content  of  the  bark 
was  produced  by  the  summer  application  of  dried  blood.  In  Sep- 
tember and  December  the  nitrogen  content  of  the  bark  on  the  sum- 


Responses  of  Apple  Trees  to  Nitrogen  Applications  15 


Table  7.  Analyses  oe  Spurs  and  Bark  From  Fertilized  and  Unfertilized 
York  Trees  in  Their  Oee  Year. 

( Percentages  of  dry  weight ) 


Dry 

weight 

Reduc’g 

sugars 

Total 

sugars 

Starch 

Ash 

K 

P 

N 

May  27,  1920 

Spurs 

Check  _ 

45.6 

1.17 

1.50 

0.92 

8.98 

0.593 

0.123 

0.773 

Blood  

44.3 

1.30 

1.56 

1.13 

8.99 

0.659 

0.125 

0.800 

Bark 

Check  

42.1 

1.38 

1.77 

0.72 

11.62 

0.601 

0.083 

0.555 

Blood  

42.5 

1.51 

1.59 

0.72 

11.80 

0.503 

0.064 

0.576 

June  24,  1920 

Spurs 

Check  

49.4 

0.92 

1.38 

2.88 

6.665 

0.516 

0.227 

0.960 

Blood  

48.1 

0.83 

1.56 

1.87 

5.818 

0.615 

0.142 

0.706 

Bark 

Check  

43.5 

0.94 



2.30 

7.845 

0.558 

0.084 

0.550 

Blood  

43.8 

1.10 

1.89 

2.00 

7.640 

0.540 

0.144 

0.721 

September  20,  1920 

Spurs 

Check  

53.9 

0.79 

1.24 

2.75 

10.38 

0.445 

0.202 

0.95 

Spring  blood  __ 

55.6 

0.61 

0.90 

2.48 

8.66 

0.394 

0.182 

0.90 

Summer  blood 

56.4 

0.61 

1.30 

1.69 

11.25 

0.461 

0.217 

1.04 

Bark 

Check  

48.1 

0.90 

1.08 

3.19 

8.87 

0.579 

0.113 

0.58 

Spring  blood  __ 

44.8 

0.70 

1.02 

2.23 

9.56 

0.603 

0.079 

0.52 

Summer  blood 

45.6 

0.79 

0.99 

2.25 

9.05 

0.658 

0.117 

0.66 

December  3,  1920 

Spurs 

Check  

44.7 

2.78 

2.79 

1.51 

8.85 

0.539 

0.233 

1.09 

Spring  blood 

46.1 

2.90 

3.00 

2.09 

10.17 

0.437 

0.214 

1.17 

Summer  blood 

46.0 

3.05 

3.30 

2.23 

9.96 

0.461 

0.240 

1.19 

Fall  nitrate 

46.8 

2.32 

2.70 

2.18 

8.82 

0.450 

0.253 

1.33 

Bark 

Check  

53.8 

4.22 

5.70 

1.91 

10.28 

0.587 

0.110 

0.72 

Spring  blood  __ 

53.4 

5.17 

5.58 

1.13 

10.14 

0.626 

0.106 

0.72 

Summer  blood 

54.6 

4.57 

4.98 

1.93 

9.84 

0.659 

0.123 

0.91 

Fall  nitrate 

54.4 

5.02 

5.22 

1.22 

9.82 

0.595 

0.159 

0.77 

March  30,  1921 

Spurs 

Check  

48.1 

1.46 

1.83 

0.81 

11.90 

0.531 

0.181 

0.85 

Spring  blood 

46.3 

1.40 

1.74 

0.77 

10.24 

0.525 

0.223 

0.92 

Summer  blood 

45.2 

1.26 

1.59 

0.45 

10.72 

0.539 

0.242 

1.01 

Fall  nitrate 

46.2 

1.19 

1.29 

0.88 

9.89 

0.438 

0.194 

1.17 

Bark 

Check 

47.3 

4.86 

5.73 

1.40 

8.82 

0.624 

0.171 

0.65 

Spring  blood  __ 

47.2 

4.48 

5.16 

2.79 

8.78 

0.666 

0.139 

0.68 

Summer  blood 

48.3 

4.73 

5.13 

2.14 

8.73 

0.654 

0.137 

0.65 

Fall  nitrate 

47.1 

4.00 

4.77 

3.58 

8.74 

0.656 

0.155 

0.70 

16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  50 


mer-fertilized  trees  was  distinctly  higher  than  that  on  the  others 
but  this  difference  had  completely  disappeared  by  the  end  of 
March,  1921. 

It  is  interesting  that  the  nitrogen  content  of  the  bark  on  the 
spring-fertilized  trees  was  apparently  increased  at  the  end  of 
June  while  the  nitrogen  content  of  the  spurs. was  much  less  at  that 
time. 


DISCUSSION 

The  analyses  reported  in  this  paper  confirm  and  extend  those 
published  in  Research  Bulletin  40  of  this  Station.  Certain  season- 
al chemical  changes  characteristic  of  bearing  and  of  non-bearing 
apple  spurs  and  certain  changes  in  bark  and  spurs  characteristic  of 
apple  trees  in  their  on  and  off  years — the  years  of  fruit  production 
and  those  of  fruit  bud  differentiation  respectively — may  be  consid- 
ered established.  Particular  physiological  processes  are  apparent- 
ly associated  with  definite  chemical  conditions  existing  in  the  spur 
at  critical  times.  Fruit  bud  differentiation,  for  example,  is  asso- 
ciated with  starch  accumulation  in  the  spur  late  in  June  and,  since 
no  exception  has  been  found  to  this  rule,  it  seems  safe  to  conclude 
that  the  conditions  which  bring  starch  accumulation  about  are 
among  the  factors  that  determine  fruit  bud  differentiation,  though 
the  existence  of  other  factors  which  are  at  times  decisive  is  un- 
questionable. In  order  to  account  for  the  known  facts  relative  to 
the  initiation  of  the  fruitful  state  it  seems  necessary  to  postulate 
two  conditions  for  fruit  bud  differentiation:  (1)  that  carbohydrate 

(or  starch)  accumulation  be  possible  and  (2)  that  no  other  factor 
such  as  nitrate,  water  or  heat  supply  be  limiting  to  the  extent  that 
vegetative  development  is  stopped  or  seriously  retarded.  These 
points  are  discussed  in  Research  Bulletin  47  of  this  Station,  where 
it  is  also  shown  that  the  conditions  determining  starch  accumula- 
tion and  fruit  bud  differentiation  are  not  always  confined  to  the 
spurs.  In  a similar  way  other  processes  are  found  to  be  associated 
with  particular  features  of  the  seasonal  chemical  picture.  Thus 
recent  investigators  have  pointed  out  that  fruit  setting  seems  to 
be  related  to  the  nitrogen  content  of  the  spur  in  May  and  vegeta- 
tive extension  is  evidently  related  to  a number  of  factors. 

The  data  presented  in  this  paper  also  show  some  of  the  effects 
of  applying  nitrogenous  fertilizers  of  various  kinds  and  at  different 
seasons  with  special  reference  to  the  chemical  composition  of  bark 
and  spurs.  These  effects  depend  primarily  on  the  condition  of  the 


Responses  of  Apple  Trees  to  Nitrogen  Applications  17 


tree  and  might  well  be  influenced  also  by  climatic  conditions.  All 
the  work  reported  here  deals  with  apple  trees  in  fairly  good  con- 
dition. The  type  of  nitrogenous  fertilizer  is  evidently  important 
under  certain  circumstances;  under  others  it  is  not.  Various  physi- 
ological processes  are  affected  more  or  less  independently  and 
there  are  indications  that  the  season  when  the  applications  are 
made  is  an  important  factor  in  determining  how  these  processes 
are  affected. 

An  intelligent  use  of  fertilizers  evidently  must  be  based  on  a 
recognition  of  the  particular  process  which  it  is  advisable  to  con- 
trol and  on  a knowledge  of  the  effects  that  applications  of  various 
kinds  and  at  different  seasons  will  have  on  this  process.  In  the 
past  when  nitrogenous  fertilizers  have  appeared  necessary,  a suit- 
able amount  has  been  determined  on  and  has  been  applied  in  the 
cheapest  or  most  readily  available  form  in  early  spring.  This  pre- 
cedure  is  inadequate  as  a panacea  and  the  facts  presented  show 
that  it  may  produce  effects  directly  opposite  to  those  desired. 
Trees  that  bear  light  crops  regularly  every  year  and  in  whose  an- 
nual yield  a material  increase  is  desired  present  a case  for  treat- 
ment as  different  from  that  of  trees  which  bear  heavy  crops  bien- 
nially and  which  it  is  desired  to  make  regular  producers  as  this  in 
turn  is  different  from  the  case  of  trees  which  bloom  profusely 
every  spring  but  set  little  or  no  crop  because  nitrogen  is  a lim- 
iting factor.  In  one  case  it  may  be  a question  of  stimulating 
general  vegetative  growth  and  vigor;  in  another  of  affecting  fruit 
bud  differentiation ; in  another  of  increasing  the  set  of  fruit.  The 
same  treatment  will  not  bring  about  the  desired  result  in  all  cases. 
Each  is  a problem  for  separate  consideration  and  each  involves 
phases  which  should  be  studied  under  a wide  variety  of  conditions. 
There  are,  of  course,  limits  to  the  effectiveness  of  nitrogenous  fer- 
tilizers, but  even  where  a requirement  for  nitrogen  can  be  estab- 
lished, the  best  method  of  application  in  one  instance  may  be,  en- 
tirely different  from  the  best  for  some  other  case.  The  effect  o( 
early  spring  applications  of  quickly  available  nitrogenous  fertilizers 
in  aiding  the  set  of  fruit  has  been  established  and  evidence  is  giv- 
en that  fruit  bud  differentiation  and  vegetative  development  also 
can  be  influenced  in  specific  directions  according  to  the  time  and 
type  of  application. 


18 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  50 


CONCLUSION 

The  chief  effects  of  spring  applications  of  nitrogenous  ferti- 
lizers to  healthy  apple  trees  are,  on  bearing  trees,  an  increased 
set  of  fruit  associated  with  a greater  nitrogen  content  in  the  spurs 
during  the  period  of  fruit  setting  and  in  non-bearing  trees  an  in- 
creased rate  of  growth.  Different  types  of  quickly  available  ni- 
trogenous fertilizers  produce  essentially  the  same  effects  though 
nitrate  of  soda  stimulated  leader  growth  on  very  young  trees  more 
than  dried  blood.  Spring  applications  of  nitrogenous  fertilizers  do 
not  favor  starch  accumulation  at  the  period  of  fruit  bud  differen- 
tiation and  consequently  they  could  not  be  expected  to  favor 
this  process.  No  effects  of  spring  applications  are  evident  in  the 
percentage  nitrogen  content  the  spring  following  the  treatment, 
though  larger  absolute  amounts  would  be  present  in  the  larger 
trees. 

The  later  in  the  season  nitrogenous  fertilizers  are  applied,  the 
greater  is  the  nitrogen  content  of  the  spurs  the  following  spring 
immediately  before  growth  begins. 


UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 

AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  51 


The  Influence  of  the  Plane  of  Nutrition 
On  the  Maintenance  Requirement 
of  Cattle 


(Publication  Authorized  November  21,  1921.) 


COLUMBIA,  MISSOURI 
FEBRUARY,  1922 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 

BOARD  OF  CONTROL. 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOtTE 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY  P.  E.  BURTON  . j.  BLANTON 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 

J.  C.  JONES,  PH.  D.,  LL.  D.,  PRESIDENT  OF  THE  UNIVERSITY 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 


STATION  STAFF 

February,  1922 


AGRICULTURAL  CHEMISTRY 

C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  A.  M. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  SievEking,  B.  S.  in  Agr. 

E.  M.  Cowan,  A.M. 

AGRICULTURAL  ENGINEERING 

J.  C.  WoolEy,  B.  S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

E.  F.  Hopkins,  Ph.  D. 

DAIRY  HUSBANDRY 
C.  Ragsdale,  B.  S.  in  Agr. 

. W.  Swett,  A.  M. 
m.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  R.  McBride,  B.  S.  in  A. 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  A.  M. 

O.  W.  Letson,  A.  M. 

B.  M.  King,  B.  S.  in  Agr. 

Alva  C.  Hill,  B.  S.  in  Agr. 

Miss  Bertha  C.  Hite,  A.B.*  Seed  Analyst. 
Miss  Pearl  Drummond,  A.  A.* 


RURAL  LIFE 
O.  R.  Johnson,  A.  M. 

S.  D.  Gromer,  A.  M. 

E.  L.  Morgan,  A.M. 

Ben  H.  Frame,  B.  S.  in  Agr. 


HORTICULTURE 

V.  R.  Gardner,  M.  S.  A. 

H.  D.  Hooker,  Jr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  M.  S. 

F.  C.  Bradford,  M.  S. 

H.  G.  Swartwout,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY 
H.  L-  Kempster,  B.  S. 

Earl  W.  Henderson,  B.S. 

SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M 

W.  A.  Albrecht,  Ph.  D. 

F.  L.  Duley,  A.M.t 

R.  R.  HudElson,  A.M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr. 

Richard  Bradfield,  A.  B. 

O.  B.  Price,  B.  S.  in  Agr. 

VETERINARY  SCIENCE 
J.  W.  Connaway,  D.  V.  S.,  M.  D. 

L.  S.  Backus,  D.  V.  M. 

O.  S.  CrislEr,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 

OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Secretary 

S.  B.  ShirkEy,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Jane  Frodsham,  Librarian. 

E.  E.  Brown,  Business  Manager. 


*In  service  of  U.  S.  Department  of  Agriculture. 
tOn  leave  of  absence. 


TABLE  OF  CONTENTS 


Introduction  - 5 

Review  of  Literature  - 5 

Experimental  Procedure  7 

Methods  Used  in  Calculating  Results  — ~ 8 

Animals  Used  in  Experiment  (Plates  I-III.)  . 9 

Average  of  Results  16 

Comparison  of  Results  19 

Comparison  of  Maintenance  Requirement  During  Summer 

and  Winter  Months  20 

Summary  and  Discussion  20 

Bibliography  21 

Original  Data  and  Calculations  in  Detail  23 

Feed  and  Weight  Record,  Table  13  24 

Dry  Matter  and  Organic  Matter  in  Feed,  Table  14  30 

Measurements  of  Experimental  Steers,  Table  15-16  40 

Measurements  of  Control  Steers,  Table  17  44 

Composition  of  Control  Steers,  Table  18  45 

Energy  Value  of  Gains  by  Periods,  Table  19  45 

Distribution  of  Control  Steers,  Table  20  46 

Comparison  of  Control  Steers  and  Experimental  Steers. 

(Figures  1-5.) 


46 


The  Influence  of  the  Plane  of  Nutrition  On  the 
Maintenance  Requirement  of  Cattle 

A.  G.  Hogan,  W.  D.  Salmon,  H.  D.  Fox 

In  1914  an  investigation*  was  begun  at  the  Missouri  Agricul- 
tural Experiment  Station  to  study  effects  of  underfeeding.  Calves 
of  beef  breeding  were  secured,  divided  into  three  groups,  and  each 
group  was  placed  on  a different  plane  of  nutrition.  Group  I was 
fed  to  grow  rapidly,  but  not  to  become  fat.  Group  II  was  placed  on 
a lower  nutritional  plane,  and  was  fed  to  gain  about  one-half  pound 
per  day.  Group  III  was  placed  on  a still  lower  plane  and  was  fed 
to  gain  about  one-third  of  a pound  per  day. 

There  were  large  differences  in  the  food  intake  of  the  three 
groups,  and  after  a considerable  amount  of  data  had  been  obtained 
it  was  decided  to  make  a study  of  the  maintenance  requirement  of 
these  steers. 

REVIEW  OF  LITERATURE 

There  has  accumulated  a considerable  mass  of  literature  con- 
cerning the  maintenance  requirement,  in  terms  of  energy,  of  animals 
as  well  as  of  man.  Much  of  this  material  has  no  direct  bearing  on 
the  problem  discussed  in  this  paper,  but  a short  historical  state- 
ment may  be  useful. 

Waters1  pointed  out  in  1908  that  if  the  ration  of  an  animal 
were  suddenly  reduced  to  a point  a little  less  than  sufficient  to 
maintain  its  weight,  there  would  be  a process  of  readjustment. 
After  a time  a stationary  live  weight  would  be  obtained  if  the 
reduction  were  not  too  severe,  and  following  that  there  might  be 
an  increase  in  weight. 

More  recent  data  from  the  Missouri  Experiment  Station2  show 
a lower  maintenance  requirement  for  animals  on  a low  plane  of 


•This  investigation  was  initiated  by  F.  B.  Mumford,  Dean  of  the  College  of  Agriculture, 
and  P.  F.  Trowbridge,  formerly  chairman  of  the  department  of  agricultural  chemistry. 
Since  September,  1918,  E.  A.  Trowbridge,  chairman  of  the  department  of  animal  husbandry, 
has  had  general  supervision  of  the  project.  This  article  was  prepared  by  A.  G.  Hogan, 
who  has  been  in  immediate  charge  since  September,  1920.  Mr.  Salmon  supervised  the  pre- 
liminary calculations,  and  calculated  the  data  for  the  summer  periods.  Mr.  Fox  made  the 
calculations  for  the  winter  periods.  A large  number  of  workers  have  contributed  to  the 
success  of  the  investigation,  but  it  does  not  seem  practicable*  to  mention  them  all  individ- 
ually. A short  article  embodying  the  essential  points  of  the  investigation  was  published  in 
the  Journal  of  Agricultural  Research,  Vol.  XXII,  p.  115. 

1Refers  to  Bibliography,  page  21. 


6 


Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


nutrition.  Three  steers  were  full-fed  until  eleven  months  old, 
then  subjected  to  a maintenance  trial.  In  order  of  economy  in 
maintenance  requirement  they  ranked  as  follows : Steer  598  first, 

Steer  596  second,  Steer  590  third.  Following  this  No.  596  was 
fed,  No.  598  was  given  one-half  productive  feed,  anT 
fourth  productive  feed.  The  steers  were  agai.  a 

maintenance  trial  in  which  they  ranked  as  follows  i,o.  590  first, 
No.  598  second,  No.  596  third.  Evidently  the  maintenance  require- 
ment closely  paralleled  the  plane  of  nutrition. 

Armsby3a  cites  the  observations  of  Zuntz  and  Hagemann  show- 
ing that  a surplus  of  feed  stimulated  muscular  activity  and  rest- 
lessness of  the  horse  to  such  an  extent  that  a ration  more  than  suf- 
ficient for  maintenance  of  this  animal  when  standing  quietly  in  its 
stall,  would  not  cause  an  increase  in  weight  under  ordinary  condi- 
tions. Experiments  with  cattle4  indicate  a similar  stimulating  ef- 
fect upon  their  muscular  movements.  Armsby,  therefore,  con- 
cludes that  at  least  a part  of  the  lower  maintenance  cost  may  come 
from  “voluntary  restriction  of  motion  on  the  part  of  the  animals 
on  a low  nutritive  plane.” 

An  experiment  by  Armsby  and  Fries5  showed  that  the  main- 
tenance requirement  of  a two-year-old  steer  was  increased  36 
percent  by  a three-month  fattening  period  in  which  the  live  weight 
was  increased  by  about  300  pounds.  The  computed  basal  metabol- 
ism per  1000  pounds  live  weight  per  day  showed  the  following  var- 
iations in  maintenance  requirement : 

In  proportion  In  proportion  to  2-3 

to  weight  power  of  live  weight 

Unfattened  4,919  cal.  5,125  cal. 

Fattened  5,275  cal.  5,943  cal. 

“The  basal  katabolism  increased  faster  than  the  body  weight  or  the 
body  surface  as  estimated  by  the  Meeh  formula.  Apparently  the 
accumulation  of  fat  tended  in  some  way  to  stimulate  the  general 
metabolism.”  Both  Kellner  and  Evvard  have  reported  data,  cited 
by  Armsby3b  showing  that  fat  steers  have  a higher  maintenance 
requirement  than  those  in  medium  condition. 

The  extensive  researches  of  F.  G.  Benedict  and  co-workers6 
in  the  field  of  human  physiology  are  of  especial  significance.  They 
have  demonstrated  that  the  basal  metabolism  of  their  subjects  was 
markedly  lower  on  a restricted  diet  than  on  a normal  diet.  In 
other  words  the  maintenance  requirement  was  lowered.  In  most 


The  Influence  of  the  Plane  of  Nutrition 


7 


cases  we  have  mentioned  it  is  impossible  to  decide  to  what  extent 
decreased  muscular  activity  accounts  for  the  lower  maintenance 
requirement  when  animals  are  on  a low  nutritive  plane. 

EXPERIMENTAL 

The  conditions  of  this  investigation  are  unique  in  one  respect, 
the  animals  were  started  on  the  project  at  weights  varying  from  154 
to  238  pounds  and  thereafter  were  kept  constantly  on  the  same 
plane  of  nutrition.  Since  the  animals  were  under  observation  for 
from  four  to  seven  years,  any  marked  or  permanent  adjustment 
to  nutritional  conditions  should  become  apparent. 

The  ideal  method  of  conducting  an  investigation  of  the  main- 
tenance requirement  of  cattle  would  provide  for  a respiration  cal- 
orimeter. Since  that  was  impossible  the  alternative  was  to  calcu- 
late the  energy  value  of  the  food  consumed,  and  correct  this  for 
the  estimated  value  of  the  gains  (or  losses)  in  body  weight.  The 
net  energy  of  the  feed  consumed  was  calculated  in  accordance 
with  procedures  developed  by  Armsby.  The  energy  values  of  the 
changes  in  body  weight  were  calculated  from  the  composition  of 
steers  that  had  been  analyzed  by  the  department  of  agricultural 
chemistry  at  this  station. 

Experimental  Animals. — Three  of  the  steers  now  under  obser- 
vation were  started  on  the  investigation  in  1914,  and  seven  others 
were  added  in  1917.  Some  of  the  more  significant  early  records 
are  condensed  in  the  following  table. 


Table  1. — Groups,  Dates  oe  Birth,  and  Breeds  oe  Animals. 


Ani- 

mal 

Group 

Date  of  birth 

Date  put  on  Exp. 

Weight 
when 
put  on 
Exp. 

Breed 

528 

I 

May  8, 

1914 

June 

11, 

1914 

157 

Hereford-high-grade 

577 

I 

March, 

1914 

Aug. 

5, 

1917 

227 

Shorthorn-grade 

571 

I 

March, 

1917 

Aug. 

5, 

1917 

158 

Hereford-grade 

579 

II 

May  2, 

1914 

May 

30, 

1914 

154 

Shorthorn-grade 

573 

II 

April, 

1917 

Aug. 

5, 

1917 

203 

Hereford-grade 

578 

II 

April, 

1917 

Aug. 

5, 

1917 

238 

Hereford-grade 

585 

III 

April  26, 

1914 

May  22, 

1914 

123 

Hereford-high-grade 

572 

III 

April, 

1917 

Aug. 

5, 

1917 

196 

Hereford-grade 

574 

III 

April, 

1917 

Aug. 

5, 

1917 

237 

Hereford-grade 

575 

III 

April, 

1917 

Aug. 

5, 

1917 

204 

Hereford-grade 

8 Missouri  Agr.  Exp.  Station  Research  Bulletin  51 

Quarters. — The  steers  had  access  to  a shed  open  to  the  south. 
Adjoining  this  shed  were  dry  lots  sloping  to  the  south,  and  having 
shade  protection. 

Rations. — The  concentrate  consisted  of  the  foil  s ' 

Corn  chop,  60  percent;  wheat  bran,  30  percent 
percent.  The  roughage  fed  from  the  beginning  of  me  experiment 
until  July  20,  1917,  was  timothy.  For  the  next  ten  days  a mixture 
of  5 parts  timothy,  3 parts  alfalfa  and  2 parts  oat  straw  was  fed. 
Following  this  the  roughage  consisted  of  a mixture  of  60  percent 
alfalfa  and  40  percent  oat  straw.  The  animals  were  fed  twice 
daily  and  had  access  to  water  at  all  times.  Salt  was  accessible 
at  feeding  time. 

Weights. — The  steers  were  weighed  each  morning,  after  feed- 
ing, but  before  watering.  The  weight  given  for  the  beginning 
of  a period  is  the  average  of  the  ten  preceding  days.  The  weight 
given  at  the  end  is  an  average  of  the  last  ten  days  of  the  period. 

Periods. — The  calculations  are  made  for  periods  of  180  days, 
with  the  exception  of  the  first  period  for  each  of  the  three  older 
steers,  which  were  as  follows:  No.  528,  130  days;  No.  579,  142 

days;  No.  585,  150  days.  In  order  that  one  period  each  year  might 
be  free  from  the  disturbing  effects  of  cold  weather,  the  year  was 
divided  into  a “summer”  and  “winter”  period.  The  summer  per- 
iods began  in  April  or  May,  and  ended  in  October.  The  winter  per- 
iods began  in  October  or  November  and  ended  in  April  or  May. 

Energy  Intake. — Our  calculations  of  the  energy  values  of  the 
feed  consumed  are  based  on  two  methods  described  by  Armsby4. 
In  one  case  the  dry  matter,  in  the  other  the  digestible  organic  nu- 
trients consumed,  was  used  to  calculate  the  net  energy  intake  of 
the  steers. 

The  method  of  calculation  based  on  dry  matter  consumed  is 
as  follows.  For  the  concentrates  the-  value  83.82  therms  per  100 
pounds  dry  matter  was  used.  This  is  the  factor  given  for  Armsby’s 
grain  mixture  No.  2*,  which  approximates  the  grain  mixture  used 
in  this  experiment.  For  timothy  hay  the  value  48.63  therms  per 
100  pounds  dry  matter  was  used.  The  factor  for  the  roughage 
mixture  used  in  the  latter  part  of  the  experiment  was  calculated 
from  the  Armsby  values,  for  alfalfa  34.10  therms,  and  for  oat  straw 

*Armsby’s  grain  mixture  No.  2 — 60  percent  corn  meal;  30  percent  crushed  oats;  10  per- 
cent O.  P.  linseed  meal. 

Our  grain  mixture— 60  percent  corn  meal;  30  percent  wheat  bran;  10  percent  O.  P. 
linseed  meal. 


The  Influence  of  the  Plane  of  Nutrition 


9 


Plate  1. — Taken  at  the  beginning  of  the  investigation. 


5?9 


10 


Missouri  Agr.  Exp.  Station  Research  Bulletin 


Plate  II. — Taken  after  being  fed  three  years  on  their  respective  nutritional 

planes. 


The  Influence  of  the  Plane  of  Nutrition 


11 


Plate  III. — Taken  after  being  fed  six  years  on  their  respective  nutritional 

planes. 


The  Influence  of  the  Plane  of  Nutrition 


13 


26.03  therms  per  100  pounds  dry  matter.  A mixture  of  60  parts 
alfalfa  and  40  parts  oat  straw  would  have  a value  of  30.87  therms 
per  100  pounds  dry  matter. 

The  values  used  are  summarized  below  in  tabular  form. 

Table  2. — Energy  Values  per  100  Pounds  Dky  Matter. 


Net  energy  values 
in  therms 


Alfalfa  hay  34.10 

Oat  straw  26.00 

Mixture,  60  percent  alfalfa,  40  percent  oat  straw  30.87 

Timothy  hay  48.63 

Grain  83.82 


The  calculations  of  the  energy  value  of  the  milk  are  also  based 
on  factors  published  by  Armsby3c.  These  are  29.01  therms  per  100 
pounds  whole  milk  (4.4  percent)  and  14.31  therms  per  100  pounds 
skim  milk  (0.2  percent).  From  these  values  factors  were  com- 
puted for  the  different  grades  of  milk  used.  The  values  used  are 
given  in  Table  3. 


Table  3. — Net  Energy  Value  oe  Milk  Used. 


Percent  fat  in  milk  Therms  net  energy  per 

hundred  pounds 


4.40 

(whole  milk) 

29.010 

3.20 

24.776 

2.70 

23.130 

1.85 

20.086 

1.20 

17.838 

0.20 

(skim  milk) 

14.310 

Thef  net  energy  intake,  based  on  digestible  organic  nutrients  con- 
sumed, was  calculated  by  the  procedure  outlined  below. 

The  Armsby  factor  for  the  metabolizable  energy  of  digestible 
organic  matter  from  roughage  is  1.588  therms  per  pound.  For 
grains  and  similar  feeds  the  factor  is  1.769  therms  per  pound. 

Armsby3d  has  also  determined  the  “average  energy  expendi- 
ture”  by  cattle  per  100  pounds  of  dry  matter  eaten.  This  is  uiven 
in  Table  4. 


14  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 
Table  4. — Energy  Expended  by  Cattle  per  100  lbs.  Dry  Matter  Consumed. 


Roughage 


Energy  expenditure 
in  therms 


Timothy  hay  

Alfalfa  hay  

Oat  straw  

Concentrate 
Grain  mixture  No.  2 


35.47 

53.03 

46.00 

51.76 


The  coefficients  of  digestibility  used  in  these  calculations  were 
derived  from  digestion  trials  conducted  under  similar  conditions  at 
this  Station.7  These  indicated  that  the  digestibility  of  the  ration 
varied  with  the  relative  amounts  of  hay  and  grain  fed.  The  fac- 
tors used  are  given  below. 


Table  5. — Digestion  Factors  eor  Organic  Matter. 


Ratio  of  1:1 

grain  to  hay 

2:3 

1:2 

1:3,  4 or  5 

1:6  or  7 

1:8,  9 or  10 

Hay  only 

Factor  .6956 

.6695 

.6434 

.6340 

.6229 

.6030 

.5832 

Inasmuch  as  the  thermal  value  of  a pound  of  organic  matter 
from  grain  differs  from  that  of  a similar  weight  of  organic  matter 
from  roughage,  the  Armsby  factors  previously  quoted  in  this  paper 
could  not  be  directly  applied  to  the  values  obtained  with  the  above 
digestion  coefficients.  Those  factors  would  not  provide  for  the 
widely  varying  proportions  of  grain  and  hay.  The  following  meth- 
od therefore  was  used  in  computing  the  energy  intake  on  the  basis 
of  digestible  organic  matter  consumed.  By  use  of  the  factors  in 
Table  5 the  wTeight  in  pounds  of  digestible  organic  matter  in  the 
mixed  ration  was  determined  for  each  period.  This  was  multi- 
plied by  1.588,  the  Armsby  factor  for  metabolizable  energy  in  a 
pound  of  digestible  organic  matter  from  hay.  The  thermal  value 
of  digstible  organic  matter  from  grain  is  1.769  however,  or  0.181 
therms  more.  Therefore  each  pound  of  digestible  organic  matter 
derived  from  grain  was  multiplied  by  0.181,  and  the  product  added 
to  the  result  obtained  by  multiplying  the  total  digestible  organic 
matter  by  1.588.  This  gave  the  total  metabolizable  energy  in  both 
the  hay  and  grain.  The  digestibility  of  the  organic  matter  of  the 


The  Influence  of  the  Plane  of  Nutrition 


15 


grain  was  estimated  by  difference.  This  ranged  closely  around  80 
percent.  The  factors  for  energy  expenditure  are  given  in  Table  4. 

It  seemed  impracticable  to  calculate  the  net  energy  of  the 
milk  consumed  on  the  basis  of  digestible  organic  matter,  so  the 
calculation  based  on  dry  matter  was  used  for  milk.  Since  the 
amount  was  small,  however,  the  method  of  calculation  would  have 
iittle  effect  on  the  final  results. 

Changes  in  Body  Weight. — In  order  to  obtain  data  concern- 
ing the  maintenance  requirement  of  these  steers,  it  is  necessary  to 
calculate  the  energy  gained  or  lost  through  changes  in  body  weight. 
Our  calculations  are  based  on  analyses  previously  made  by  the 
department  of  agricultural  chemistry*,  University  of  Missouri. 
Control  animals  were  selected  from  those  on  which  analyses  were 
available,  on  the  basis  of  similar  weights  and  measurements,  and 
when  possible  of  similar  ages,  daily  gains  and  daily  consumption  of 
dry  matter.  In  some  cases  suitable  check  animals  were  not  avail- 
able, and  the  composition  of  steers  for  those  periods  was  esti- 
mated by  interpolation,  using  data  of  the  preceding  and  succeeding 
periods.  Using  these  assumed  values  for  the  composition  of  the 
steers  during  the  different  periods,  the  gain  in  protein  and  fat 
was  readily  calculated.  The  thermal  equivalent  of  the  protein  and 
fat  gained  was  calculated  from  data  obtained  by  other  investigators. 
Armsby3e  quotes  data,  computed  by  Kohler,  giving  the  value  5.6776 
calories  per  gram  or  2.5753  therms  per  pound,  for  protein  of  mus- 
cular tissue  of  cattle. 

Fries8  gives  an  average  value  of  9.4889  calories  per  gram  of 
beef  fat,  or  4.3048  therms  per  pound. 

Since  no  suitable  control  animal  was  available  for  the  last  two 
periods  of  Steer  528,  the  gains  for  these  two  periods  in  terms  of 
protein  and  fat  were  not  calculated,  and  the  energy  value  of  a pound 
gain  was  assumed  to  be  3,000  therms  per  pound.  This  is  the 
value  given  by  Armsby3f  for  animals  of  apparently  similar  condi- 
tion. 

The  values  we  have  used,  also  those  published  by  Armsby 
are  given  in  Table  6.  Armsby’s  values  are  consistently  higher,  as 
is  to  be  expected.  Our  animals  were  thin,  and  contained  less  than 
the  usual  amount  of  fat  in  the  gain. 

In  calculating  the  maintenance  requirements  per  1,000  pounds 
live  weight,  Moulton’s9  formula  was  used.  He  has  shown  that 


•These  have  not  as  yet  been  published. 


16  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  6. — Energy  Value  per  Pound  Gain. 


Approximate 

age 

Group 

Armsby’s 

> Values 

I 

II 

III 

Age 

Energy 

months 

therms 

therms 

therms 

months 

therms 

6 

.95575 

.95575 

.8343 

1 

1.170 

18 

1.0918 

1.0583 

.9445 

2-3 

1.374 

36 

1.7136 

1.1608 

1.0548 

5-6 

1.680 

54 

2.1993 

1.4104 

1.1013 

11-12 

2.292 

66 

2.50 

1.5352 

1.4790 

18-24 

3.000 

78 

3.00 

1.660 

1.6490 

the  surface  areas  of  thin  cattle  are  proportional  to  the  five-eighths 
power  of  the  live  weight. 

The  average  maintenance  requirement  of  the  three  groups  is 
given  in  Tables  7 to  10  inclusive.  The  summer  and  winter  periods 
are  given  separately,  and  each  has  been  calculated  by  two  methods. 
One  is  based  on  dry  matter  consumed,  the  other  on  digestible  or- 
ganic nutrients  consumed. 

In  calculating  the  maintenance  requirement  on  the  basis  of  di- 
gestible organic  matter  consumed,  digestion  coefficients  were  used 
that  had  . been  obtained  at  this  station  under  similar  conditions. 
This  method  is  probably  more  accurate  than  the  one  based  on  dry 
matter  consumed,  and  in  this  case  gives  a result  somewhat  higher. 

During  the  summer  months  there  were  four  periods  in  all  in 
which  losses  in  live  weight  occurred.  In  calculating  averages  those 
periods  were  omitted,  as  the  results  are  low.  It  is  possible  that 


Table  7. — Average  Daily  Maintenance  Requirement  During  Summer 
Periods  as  Calculated  from  Dry  Matter  Consumed. 


Steer  number 

Therms  net  energy  per  1000  pounds 
based  on  5-8  power  of  live  weight 

Group  I 

Group  II 

Group  III 

528 — Average  of  6 periods 

5.870 

5.280 

5.073 

577 — Average  of  3 periods 

571 — Average  of  3 periods 

579 — Average  of  5 periods 

4.920 

3.830 

4.409 

578 — Average  of  3 periods 

573 — Average  of  3 periods 

585 — Average  of  5 periods 

4.221 

4.041 

4.302 

3.256 

3.921 

575 — Average  of  3 periods 

574 — Average  of  3 periods 

572 — Average  of  3 periods 

Average  of  each  group 

5.523 

4.483 

The  Influence  the  Plane  of  Nutrition 


17 


Table  8. — Average  Daily  Maintenance  Requirement  During  W inter  Months 
as  Calculated  from  Dry  Matter  Consumed. 


Steer  number 

Therms  net  energy  per  1000  pounds 
based  on  5-8  power  of  live  weight 

Group  I 

Group  II 

Group  III 

528 — Average  of  6 periods 

5.909 

577 — Average  of  3 periods 

5.530 

571 — Average  of  3 periods  

5.450 

579 — Average  of  6 periods 

4.647 

578 — Average  of  3 periods 

3.730 

573 — Average  of  3 periods 

4.753 

585 — Average  of  6 periods 

• 4.366 

575 — Average  of  3 periods 

4.260 

574 — Average  of  3 periods 

4.673 

572 — Average  of  3 periods  

3.157 

Average  of  each  group 

5.770 

4.444 

4.164 

they  are  correct,  but  the  apparently  diminished  requirement  may 
be  due  to  an  incorrect  assumption  as  to  the  energy  value  of  the 
loss  in  weight.  One  steer,  No.  585,  had  a navel  infection  during 
the  first  summer  period,  accompanied  by  a very  high  maintenance 
requirement.  This  period  also  was  discarded  in  calculating  aver- 
ages. 

There  is  a close  parallel  between  the  intake  of  net  energy  and 
the  maintenance  requirement  of  the  animal.  The  record  of  Steer 
574  for  the  summer  periods  illustrates  that  tendency.  For  the  first 
period  the  average  daily  intake  of  net  energy  was  3.884  therms  per 
1,000  pounds,  based  on  the  five-eighths  power  of  the  live  weight, 
and  the  maintenance  requirement  was  3.818  therms.  For  the  second 
period  the  energy  intake  was  increased  to  5.783  therms,  and  the 


Table  9. — Average  Daily  Maintenance  Rrquirement  During  Summer  Months 
as  Calculated  from  Digestible  Organic  Matter  Consumed. 


Steer  number 

Therms  net  energy  per  1000  pounds 
based  on  5-8  power  of  live  weight 

Group  I 

Group  II 

Group  III 

528 — Average  of  6 periods 

6.261 

577 — Average  of  3 periods 

5.412 

571 — Average  of  3 periods 

5. 174 

579 — Average  of  5 periods 

5.260 

578 — Average  of  3 periods 

4.192 

573 — Average  of  3 periods 

4.893 

585 — Average  of  5 periods 

4.725 

575 — Average  of  3 periods 

4.454 

574 — Average  of  3 periods 

4.591 

572 — Average  of  3 periods 

3.649 

Average  of  each  group 

5.777 

4.869 

4.408 

18  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  10.— Average  Daily  Maintenance  Requirement  During  Winter 
Months  as  Calculated  from  Digestible  Organic  Matter  Consumed 


Steer  number 

Therms  net  energy  per  1000  pounds 
based  on  5-8  power  of  live  weight 

Group  I 

Group  II 

Group  III 

528 

5.965 

5.713 

5.553 

577 

571 

579 

5.071 

4.494 

5.513 

578 

573 

585 * 

4.625 

4.429 

5.290 

3.754 

575 

574 

572 

Average  of  each  group 

5.799 

5.037 

4.869 

maintenance  requirement  increased  to  5.119  therms.  In  the  third 
period  the  energy  intake  was  5.253  therms,  and  the  maintenance 
requirement  was  4.836  therms. 

In  comparing  the  maintenance  requirements  of  the  three  groups 
it  should  be  kept  in  mind  that  Group  I does  not  represent  a high 
plane  of  nutrition.  The  aim  was  to  secure  maximum  growth  with 
no  considerable  fattening.  Their  maintenance  requirements  as 
computed  in  this  paper  correspond  closely  to  the  average  of  22 
respiration  experiments  by  Armsby  and  Fries,  and  seven  by  Kellner 
on  cattle  in  medium  condition.  A comparison  of  our  results  (com- 
puted on  the  basis  of  digestible  organic  matter  consumed)  and  of 
those  obtained  by  other  investigators  is  given  in  Table  11. 

A few  facts  not  shown  by  the  data  seem  worthy  of  record. 
Although  some  of  the  steers  were  receiving  a very  scanty  ration, 
they  apparently  did  not  have  an  unusual  desire  for  food,  and  some 
care  was  necessary  to  prevent  their  “getting  off  feed.”  This  is 
especially  true  of  the  roughage,  for  it  was  impossible  to  induce 
them  to  consume  a large  quantity  of  hay.  Any  increase  in  the 
grain  ration  had  to  be  very  gradual. 

The  dentition  of  these  steers  was  apparently  the  same  as  for 
normal  animals,  as  regards  age.  So  far  as  could  be  determined  by 
observation,  the  temporary  teeth  were  lost  at  the  normal  age. 

Influence  of  Age. — The  ages  represented  in  this  experiment 
vary  from  30  days  for  some  of  the  calves  at  the  beginning  of  the 
first  summer  period  to  more  than  six  years  at  the  close  of  the 
seventh  period.  Apparently  there  was  no  relation  between  the  age 


The  Influence  of  the  Plane  of  Nutrition 


19 


Table  11. — Daily  Maintenance  Requirements  of  Cattle  — Net  Energy 


No.  of 
Exper- 
iments 

Investigator 

Condition 

of 

animal 

Therms  per  1000  lbs.  live  wt. 

Maximum 

Minimum 

Average 

Respiration  Exp’s. 

22 

Armsby  and  Fries  (3b) 

Medium 

7.430 

4.723 

5.995 

r.  7 

Kellner  (3b) 

Medium 

6.780 

4.921 

5.742 

Kellner  (3b) 

Fat 

8.871 

7.319 

7.946 

Live  Wt.  Exp’s. 

10 

Armsby  (3b) 

Thin 

7.044 

6.136 

6.505 

3 

Armsby  (3b) 

Thin 

6.039 

4.713 

5.423 

6 

Haecker  (3b) 

Medium 

5.676 

4.662 

5.021 

3 

Eward  (3b) 

Medium 

7.079 

5.841 

6.173 

7 

Eckles  (3b) 

Medium 

7.079 

5.841 

6.173 

1 

Shirky  (7) 

Medium  a 

7.732 

2 

Shirky  (7) 

Thin  b 

5.0959 

4.953 

5.0245 

3 

Our  results,  summer  periods 

Group  I 

7.380 

4.915 

5.777 

3 

Our  results,  summer  periods 

Group  II 

5.724 

3.809 

4.869 

4 

Our  results,  summer  periods 

Group  III 

5.217 

3.276 

4.408 

3 

Our  results,  winter  periods . 

Group  I 

7.431 

4.314 

5.799 

3 

Our  results,  winter  periods . 

Group  II 

7.598 

3.246 

5.037 

4 

Our  results,  winter  periods . 

Group  III 

5.574 

3.475 

4.869 

a Corresponds  to  Group  I animals  this  experiment. 
b Corresponds  to  Group  II  of  this  experiment. 


and  the  maintenance  requirement  of  these  animals.  Some  of  the 
steers  showed  a gradual  decrease  in  the  maintenance  cost  from  the 
beginning  to  the  end  of  the  experiment.  In  such  cases  it  was 
found  that  the  energy  intake  per  1,000  pounds  had  also  decreased. 
On  the  other  hand,  steers  with  an  increasing  energy  intake  showed 
an  increased  maintenance  requirement.  Maintenance  trials  on 
young  animals  usually  give  higher  results  than  have  been  obtained 
with  mature  animals,  but  if  age  does  influence  the  maintenance 
requirement  the  effect  is  too  slight  to  be  shown  in  a live  weight  ex- 
periment of  this  kind. 

Influence  of  Season. — The  maintenance  requirement  of  the 
steers  in  Groups  I and  II  is  slightly  higher  during  the  winter,  as 
compared  to  the  summer  months.  The  animals  in  Group  III  how- 
ever required  considerably  more  energy  for  maintenance  during  the 
winter  periods  than  they  did  in  the  summer  periods.  Presumably 
the  energy  expenditure  incident  to  the  greater  consumption  of  feed 
by  the  steers  of  Groups  I and  II  is  sufficiently  great  to  make  un- 
necessary the  oxidation  of  a large  quantity  of  additional  nutrients 
during  the  winter  months' in  order  to  maintain  the  body  tempera- 
ture. This  is  not  the  case  with  the  steers  on  a lower  nutritional 
plane,  and  so  during  periods  of  prevailingly  low  temperatures  they 


20  Missouri  Agricultural  Experiment  Station  Bulletin  51 


must  oxidize  a larger  amount  of  material  in  order  to  counteract  the 
more  rapid  loss  of  heat  from  the  body  surface.  The  contrast  be- 
tween the  two  seasons  is  shown  in  Table  12. 

Table  12— Daily  Maintenance  Requirement  in  Therms  oe  Cattle  During 
Summer  and  Winter  Months. 


During  Summer  and  Winter  Months 
Calculated  on  basis  of  digestible  organic  matter  consumed 


Group  I 

Group  II 

Group  III 

Summer 

5.777 

4.869 

4.408 

Winter 

5.779  5.037 

Calculated  on  basis  of  dry  matter  consumed 

4.869 

Summer 

5.523 

4.483 

3.921 

Winter 

5.770  4.444 

Average  of  results  obtained  by  the  two  methods 

4.164 

Summer 

5.650 

4.676 

4.165 

Winter 

5.775 

4.741 

4.517 

SUMMARY  AND  DISCUSSION 

There  is  a close  relation  between  the  amount  of  net  energy 
consumed  and  the  maintenance  requirement.  Periods  of  high  en- 
ergy intake  were  periods  of  high  maintenance  cost,  while  periods 
of  low  energy  intake  were  accompanied  by  a lowered  maintenance 
requirement. 

The  averages  of  all  periods  show  the  following  daily  mainten- 
ance requirements  per  1,000  pounds  live  weight,  in  terms  of  net 
energy.  Summer  months : Group  I,  (high  plane)  5.650  therms ; 

Group  II,  (medium  plane)  4.676  therms;  Group  III,  (low  plane) 
4.165  therms.  Winter  months:  Group  I,  5.775  therms;  Group  II, 
4.741  therms;  Group  III,  4.517  therms. 

The  maintenance  requirement  of  Group  I is  about  20  percent 
higher  than  that  of  Group  II,  and  about  30  percent  higher  than  that 
of  Group  III. 

If  there  is  a definite  relation  between  the  age  of  animals  and 
their  maintenance  requirements,  it  was  obscured  in  this  investi- 
gation by  variations  in  the  food  intake. 

The  maintenance  requirement  of  these  animals  is  higher  in  the 
winter  than  in  the  summer. 


The  Influence  of  the  Plane  of  Nutrition 


21 


BIBLIOGRAPHY 


1.  Waters,  H.  J. 

1908 — Capacity  of  Animals  to  Grow  Under  Adverse  Conditions.  Proc. 
Soc.  Promotion  Agr.  Science  29th  Annual  Meeting,  p.  71. 

2.  Trowbridge,  P.  F.,  Moulton,  C.  R.,  Haigh,  L.  D. 

1915 — The  Maintenance  Requirement  of  Cattle,  Missouri  Agricultural 
Experiment  Station,  Research  Bulletin  18. 

3.  Armsby,  H.  P. 

1917 — The  Nutrition  of  Farm  Animals.  New  York,  The  MacMillan 
Company. 

(a)  Page  306  (c)  Page  719  (e)  Page  54 

(b)  Page  291  (d)  Page  652  (f)  Page  400 

4.  Armsby,  H.  P.,  Fries,  J.  A. 

1915 —  Net  Energy  of  deeding  Stuffs  for  Cattle.  Jour.  Agr.  Research 
Vol.  3,  p.  435. 

5.  Armsby,  H.  P.,  Fries,  J.  A. 

1917 — Influence  of  Degree  of  Fatness  of  Cattle  Upon  Their  Utiliza- 
tion of  Feed.  Jour.  Agr.  Research,  Vol.  11,  p.  451. 

6.  Benedict,  F.  G.,  Miles,  W.  R.,  Roth,  Paul,  Smith,  H.  M. 

Human  Vitality  and  Efficiency  under  Prolonged  Restricted  Diet. 
Carnegie  Institution  of  Washington,  Publication  No.  280. 

7.  Shirky,  S.  B. 

1919 — The  Extent  to  Which  Growth  Retarded  During  the  Early  Life 
of  the  Animal  Can  Be  Regained.  University  of  Missouri,  Thesis  for 
the  degree,  Master  of  Arts. 

8.  Fries,  J.  A. 

1907 — Investigations  in  the  Use  of  the  Bomb  Calorimeter.  U.  S.  Dept, 
of  Agriculture,  Bur.  Animal  Industry,  Bulletin  94,  p.  13. 

9.  Moulton,  C.  R. 

1916 —  Units  of  Reference  for  Basal  Metabolism  and  Their  Interre- 
lations. Jour.  Biol.  Chem.,  Vol.  24,  p.  299. 


ORIGINAL  DATA  AND  CALCULATIONS 
IN  DETAIL 


24  Missouri  Agricultural  Experiment  Station  Bulletin  51 


Table  13. — Weight  in  Pounds  of  Animals,  and  of  Feed  Consumed  by  Thirty 

Day  Periods 


Date 
beginning 
of  period 

Period 

No. 

Live0 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Date 
beginning 
of  period 

Period 

No. 

Live0 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Steer  585 

585  (Cont. 

) 

5-22-14 

1 

143.1 

8.75 

321 

8-  5-17 

40 

495 

15.0 

220.5 

6-21-14 

2 

155 

5.00 

15.5 

317.5 

9-  4-17 

41 

498 

37.0 

225.0 

7-21-14 

3 

173 

59.5 

300 

10-  4-17 

42 

512 

70.0 

225.0 

8-20-14 

4 

188 

90.0 

300 

11-  3-17 

43 

529 

67.0 

232.0 

9-19-14 

5 

194 

104.0 

299.8 

12-  3-17 

44 

532 

72.5 

240.0 

10-18-14 

6 

200 

120.0 

290.0 

1-  2-18 

45 

554 

76.0 

240.0 

11-18-14 

7 

192 

140.0 

100.0 

2-  1-18 

46 

576 

36.5 

240.0 

12-18-14 

8 

198 

150  0 

3-  3-]8 

47 

586 

50.0 

236.0 

1-17-15 

9 

193 

150  0 

4-  2-18 

48 

576 

225.0 

2-16-15 

10 

194 

150  0 

5-  2-18 

49 

596 

224.5 

3-18-15 

11 

218 

150  0 

6-  1-18 

50 

578 

240.0 

4-17-15 

12 

223  ' 

150.0 

7-  1-18 

51 

568 

240.0 

5-17-15 

13 

229 

150.0 

7-31-18 

52 

554 

240.0 

6-16-15 

14 

233 

155.0 

8-30-18 

53 

542 

253.0 

7-17-15 

15 

235 

11  75 

153  5 

9_29-] 8 

54 

532 

255.0 

8-16-15 

16 

251 

36.7 

163.5 

10-29-18 

55 

506 

263.5 

9-15-15 

17 

256 

47.0 

150.5 

1 1-28-18 

56 

532 

278.0 

10-14-15 

18 

272 

60.0 

154  0 

12-28-18 

57 

558 

328.5 

11-14-15 

19 

290 

60.0 

180.0 

1-27-19 

58 

562 

317.0 

12-14-15 

20 

303 

60.0 

180.0 

2-26-19 

59 

591 

330.0 

1-13-16 

21 

312 

60.0 

180.0 

3-28-19 

60 

603 

330.0 

2-12-16 

22 

337 

56.5 

180.0 

4-27-1 9 

61 

604 

317.5 

3-13-16 

23 

345 

34.0 

180.0 

5-27-19 

62 

614 

326.5 

4-12-16 

24 

356 

30.0 

180.0 

6-26-19 

63 

622 

335.0 

5-12-16 

25 

360 

30.0 

180.0 

7-26-19 

64 

643 

360.0 

6-11-16 

26 

361 

31 .5 

180.0 

8-25-19 

65 

630 

360.0 

7-11-16 

27 

370 

45.0 

180.0 

9-24-19 

66 

645 

31.4 

316.5 

8-10-16 

28 

387 

45.0 

180.0 

10-24-19 

67 

653 

60.0 

333.0 

9-  9-16 

29 

386 

42.5 

170.0 

11-23-19 

68 

688 

58.0 

257.5 

10-  9-16 

30 

403 

45.0 

187.5 

12-23-19 

69 

682 

60.0 

334.0 

11-  8-16 

31 

413 

45.0 

201.0 

1-22-20 

70 

716 

60.0 

360.0 

12-  8-16 

32 

419 

51.0 

211.5 

2-21-20 

71 

741 

60.0 

360.0 

1-  7-17 

33 

437 

75.0 

225.0 

3-22-20 

72 

763 

60.0 

360.0 

2-  6-17 

34 

465 

66.0 

218.5 

4-21-20 

73 

785 

60.0 

360.0 

3-  8-17 

35 

484 

55.0 

210.0 

5-21-20 

74 

825 

60.0 

359.0 

4-  7-17 

36 

482 

38.0 

210.0 

6-20-20 

75 

850 

60.0 

360.0 

5-  7-17 

37 

496 

30.0 

210.0 

7-20-20 

76 

858 

60.0 

359.0 

6-  6-17 

38 

499 

25.0 

210.5 

8-19-20 

77 

878 

60.0 

360.0 

7-  6-17 

39 

504 

15.0 

208.5 

9-18-20 

78 

885 

60.0 

360.0 

Steer  579 

579  (Cont. 

) 

5-30-14 

1 

154.0 

2.55 

226 

8-  5-17 

40 

709 

75.0 

225.0 

6-21-14 

2 

183.9 

7.3 

14.5 

374 

9-  4-17 

41 

707 

86.0 

225.0 

7-21-14 

3 

204.6 

24.0 

47.0 

420 

10-  4-17 

42 

725 

141.5 

231.0 

8-20-14 

4 

240.0 

30.0 

60.5 

208 

11-  3-17 

43 

744 

136.5 

247.5 

9-19-14 

5 

270 

37.0 

74.0 

420 

12-  3-17 

44 

758 

147.0 

251.5 

10-18-14 

6 

304 

45.0 

90.0 

420 

1-  2-18 

45 

789 

164.5 

253.0 

11-18-14 

7 

305 

45.0 

103.0 

140 

2-  1-18 

46 

820 

120.0 

255.0 

12-18-14 

8 

316 

45.0 

120.0 

3-  3-18 

47 

822 

67.0 

255.0 

1-17-15 

9 

321 

45.0 

120.0 

4-  2-18 

48 

831 

73.5 

255.5 

2-16-15 

10 

322 

45.0 

122.0 

5-  2-18 

49 

851 

22.5 

292.5 

3-18-15 

11 

335 

45.0 

120.0 

6-  1-18 

50 

834.5 

309.0 

“Average  weight  of  last  ten  days  of  period. 


The  Influence  of  the  Plane  of  Nutrition 


25 


Table  13  (Continued). — Weight  in  Pounds  of  Animals  and  of  Feed  Consumed 
by  Thirty  Day  Periods 


Date 

beginning 
of  period 

Period 

No. 

Live0 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Date 

beginning 
of  period 

Period 

No. 

Live0 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Steer  579 

(Cont.) 

Steer  579 

(Cont.) 

4_17_15 

12 

334 

120.0 

7-  1-18 

51 

821 

315.0 

5-17-15 

13 

344 

45.0 

120.0 

7-31-18 

52 

786 

315.0 

6-16-15 

14 

344 

50.0 

120.0 

8-30-18 

53 

788 

26.0 

330.0 

7 17-15 

15 

351 

71  5 

143  0 

9-29-18 

54 

781 

2.0 

339.0 

8-16-15 

16 

379 

76  25 

178  0 

10-29-18 

55 

705 

253.0 

9-15-15 

17 

399 

70.0 

180  0 

11-28-18 

56 

730 

304.5 

10-14-15 

18 

409 

60.0 

180  0 

12-28-18 

57 

750 

330.0 

11-14-15 

19 

419 

64.5 

184  5 

1-27-19 

58 

750 

351.5 

12-14-15 

20 

430 

76.5 

194.0 

2-26-19 

59 

780 

360.0 

1-13-16 

21 

434 

75.0 

195  0 

3-28-1 9 

60 

788 

360.0 

2-12-16 

22 

453 

75  0 

195  0 

4-27-19 

61 

808 

357.0 

3-13-16 

23 

469 

75.0 

195.0 

5-27-19 

62 

729 

287.0 

4-12-16 

24 

489 

75.0 

195.0 

6-26-19 

63 

776 

326.0 

5-12-16 

25 

495 

74.0 

194.0 

7-26-19 

64 

799 

359.0 

6-11-16 

26 

505 

68.0 

180.0 

8-25-19 

65 

776 

360.0 

7-11-16 

27 

521 

75.0 

180.0 

9-24-19 

66 

766 

8.5 

360.0 

8-10-16 

28 

536 

75.0 

180.0 

10-24-19 

67 

771 

59.0 

360.0 

9-  9-16 

29 

552 

75.0 

180.0 

11-25-19 

68 

782 

60.0 

360.0 

10-  9-16 

30 

557 

75.0 

187.5 

12-23-19 

69 

797 

60.0 

329.5 

11-  8-16 

31 

565 

81.0 

189.0 

1-22-20 

70 

823 

60.0 

360.0 

12-  8-16 

32 

560 

90.0 

200.0 

2-21-20 

71 

843 

60.0 

360.0 

1-  7-17 

33 

582 

90.0 

216.0 

3-22-20 

72 

855 

60.0 

360.0 

2-  6-17 

34 

601 

93.0 

211.0 

4-21-20 

73 

857 

60.0 

360.0 

3-  8-17 

35 

632 

105.0 

225.0 

5-21-20 

74 

866 

59.0 

328.0 

4-  7-17 

36 

653 

95.5 

224.5 

6-20-20 

75 

893 

60.0 

360.0 

5-  7-17 

37 

674 

94.0 

229.5 

7-20-20 

76 

910 

60.0 

358.0 

6-  6-17 

38 

701 

91.5 

224.5 

8-19-20 

77 

907 

60.0 

360.0 

7-  6-17 

39 

708 

75.0 

225.0 

9-19-20 

78 

918 

60.0 

360.0 

Steer  528 

528  (Cont. 

) 

6-11-14 

1 

179.0 

2.5 

124.9 

8-  5-17 

40 

1008 

188.0 

227.0 

6-21-14 

2 

202.8 

14.0 

14.0 

489 

9-  4-17 

41 

1029 

196.0 

238.0 

7-21-14 

3 

252.0 

44.0 

53.5 

582 

10-  4-17 

42 

1043 

209.0 

231.0 

8-20-14 

4 

303.8 

69.0 

75.0 

600 

11-  3-17 

43 

1070 

218.0 

240.0 

9-19-14 

5 

363.8 

110.5 

101.5 

600 

12-  3-17 

44 

1076 

225.0 

240.0 

10-18-14 

6 

425.1 

120.0 

120.0 

600 

1-  2-18 

45 

1080 

225.0 

240.0 

11-18-14 

7 

426.8 

105.2 

120.0 

176 

2-  1-18 

46 

1178 

225.0 

240.0 

12-18-14 

8 

439.4 

120.0 

120.0 

3-  3-18 

47 

1147 

225.0 

240.0 

1-17-15 

9 

457.2 

120.0 

120.0 

4-  2-18 

48 

1163 

224.5 

240.5 

2-16-15 

10 

460.6 

120.0 

119.0 

5-  2-18 

49 

1184 

150.0 

293.5 

3-18-15 

11 

482 

120.0 

120.0 

6-  1-18 

50 

1177 

97.0 

318.0 

4-17-15 

12 

497 

120.0 

120.0 

7-  1-18 

51 

1159 

28.0 

330.0 

5-17-15 

13 

507 

120.0 

120.0 

7-31-18 

52 

1139 

30.0 

330.0 

6-16-15 

14 

517 

120.0 

130.0 

8-30-18 

53 

1122 

30.0 

330.0 

7-17-15 

15 

527 

120.0 

150.0 

9-29-18 

54 

1107 

30.0 

336.0 

8-16-15 

16 

559 

145.7 

152.0 

10-29-18 

55 

1103 

87.0 

333.0 

9-15-15 

17 

586 

150.0 

180.0 

11-28-18 

56 

1122 

120 

353.5 

10-14-15 

18 

620 

150.0 

180.0 

12-28-18 

57 

1155 

120.0 

357.5 

11-14-15 

19 

629 

155.0 

184.5 

1-27-19 

58 

1158 

120.0 

360.0 

12-14-15 

20 

651 

165.0 

195.0 

2-26-19 

59 

1172 

150.5 

360.0 

1-13-16 

21 

659 

165.0 

195.0 

3-28-19 

60 

121 1 

150.0 

360.0 

2-12-16 

22 

678 

165.0 

195.0 

4-27-19 

61 

1231 

150.0 

360.0 

“Average  weight  of  last  ten  days  of  period. 


26  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  13  (Continued). — Weight  in  Pounds  of  Animals  and  of  Feed  Consumed 
by  Thirty  Day  Periods 


Date 

beginning 
of  period 

Period 

No. 

Live0 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Date 
beginning 
of  period 

Period 

No. 

Live0 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Steer  528 

(Cont.) 

Steer  628 

(Cont.) 

3-13-16 

23 

699 

165.0 

195.0 

5-27-19 

62 

1258 

150.0 

360.0 

4-12-16 

24 

727 

165.0 

195.0 

6-26-19 

63 

1244 

149.0 

357.0 

5-12-16 

25 

745 

165.0 

195.0 

7-26-19 

64 

1267 

150.0 

360.0 

6-11-16 

26 

768 

165.0 

194.5 

8-25-19 

65 

1278 

150.0 

360.0 

7-11-16 

27 

785 

165.0 

180.0 

9-24-19 

66 

1278 

150.0 

360.0 

8-10-16 

28 

805 

165.0 

180.5 

10-24-19 

67 

1274 

150.0 

360.0 

9-  9-16 

29 

837 

179.5 

189.0 

11-23-19 

68 

1285 

148.5 

360.0 

10-  9-16 

30 

869 

180.0 

202.5 

12-23-19 

69 

1304 

150.0 

360.0 

11-  8-16 

31 

881 

180.0 

211.0 

1-22-20 

70 

1314 

150.0 

360.0 

12-  8-16 

32 

878 

180.0 

214.5 

2-21-20 

71 

1333 

150.0 

360.0 

1-  7-17 

33 

887 

180.0 

226.0 

3-22-20 

72 

1342 

150.0 

360.0 

2-  6-17 

34 

915 

180.0 

225.0 

4-21-20 

73 

1341 

150.0 

360.0 

3-  8-17 

35 

933 

180.0 

225.5 

5-21-20 

74 

1355 

150.0 

360.0 

4-  7-17 

36 

942 

180.0 

225.0 

6-20-20 

75 

1367 

150.0 

360.0 

5-  7-17 

37 

964 

190.0 

228.5 

7-20-20 

76 

1369 

150.0 

360.0 

6-  6-17 

38 

986 

180.0 

225.0 

8-19-20 

77 

1381 

147.0 

348.0 

7-  6-17 

39 

990 

180.0 

225.0 

9-19-20 

78 

1401 

150.0 

360.0 

Date 

beginning 
of  period 


Period 

No. 


Live0 

weight 

Pounds 


Grain 

Pounds 


Hay 

Pounds 


Milk 

Pounds 


Period 

No. 


Live0 

weight 

Pounds 


Grain  Hay 
Pounds  Pounds 


Milk 

Pounds 


Steer  578 

8-  5-17 

1 

9-  4-17 

2 

10-  4-17 

3 

11-  8-17 

4 

12-  3-17 

5 

1-  2-18 

6 

2-  1-18 

7 

3-  3-18 

8 

4-  2-18 

9 

5-  2-18 

10 

6-  1-18 

11 

7-  1-18 

12 

7-31-18 

13 

8-30-18 

14 

9-29-18 

15 

10-29-18 

16 

11-28-18 

17 

12-28-18 

18 

1-27-19 

19 

2-26-19 

20 

3-28-19 

21 

4-27-19 

22 

5-27-19 

23 

6-26-19 

24 

7-26-19 

25 

8-25-19 

26 

9-24-19 

27 

252 

268 

279 

314 

317 

331 

365 

383 

378 

411 

418 

413 

402 

393 

383 

372 

375 

395 

409 

432 

438 

455 

458 

457 

466 

490 

496 


25.0 

99.5 

152.5 

44.3 

111.0 

216.4 

72.2 

130.8 

54.5 

90.0 

140.4 

76.1 

194.0 

81.0 

164.5 

81.6 

165.0 

29.0 

164.0 

21.5 

165.5 

10.0 

192.0 

196.0 

195.0 

195.0 

208.0 

210.0 

2.0 

210.0 

233.5 

226.5 

240.0 

240.0 

240.0 

240.0 

241.5 

245.0 

17.0 

269.0 

30.0 

270.0 

30.5 

270.0 

Steer 

577 

1 

249 

2 

272 

3 

288 

4 

317 

5 

339 

6 

361 

7 

404 

8 

441 

9 

472 

10 

504 

11 

524 

12 

538 

13 

555 

14 

560 

15 

578 

16 

599 

17 

617 

18 

640 

19 

671 

20 

690 

21 

730 

22 

750 

23 

764 

24 

778 

25 

813 

26 

820 

27 

810 

23 

.5 

79, 

.4 

33 

.0 

104 

.1 

71 

.1 

118. 

.2 

94 

.9 

140. 

.4 

105. 

.0 

194. 

4 

105 

.5 

164. 

.5 

107. 

.5 

165. 

.0 

120. 

.0 

165. 

.0 

119 

.5 

165. 

.5 

91 

.5 

195. 

.0 

90. 

.0 

198. 

.5 

90 

.0 

210. 

,0 

90 

.0 

210 

.0 

90. 

.0 

223. 

.0 

90, 

.0 

225. 

.0 

113. 

.0 

211 

.5 

120. 

.0 

210. 

0 

120. 

0 

210. 

0 

120 

.0 

231. 

0 

120 

.0 

240. 

0 

120. 

.0 

240. 

0 

119 

.0 

228. 

,5 

120 

.0 

240. 

.0 

120 

.0 

240. 

0 

119 

.0 

240. 

.0 

120 

.0 

239. 

.0 

107 

.5 

226. 

.5 

’Average  weight  of  last  ten  days  of  period. 


The  Influence  of  the  Plane  of  Nutrition 


27 


Table  13  (Continued). — Weight  in  Pounds  of  Animals  and  of  Feed  Consumed 

by  Thirty  Day  Periods 


Date 
beginning 
of  period 

Period 

No. 

Live" 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Period 

No. 

Live" 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Steer  578 

(Cont.) 

Steer 

577  (Co 

nt.) 

10-24-19 

28 

512 

30.0 

270.0 

28 

833 

120.0 

258.0 

1 1-23-19 

29 

519 

30  0 

270.0 

29 

855 

120.0 

270.0 

12  23  IQ 

30 

541 

30  0 

270  0 

30 

875 

120.0 

270.0 

1-22-20 

31 

555 

30  0 

270  0 

31 

905 

120.0 

270.0 

2-21-20 

32 

580 

30.0 

270.0 

32 

915 

120.0 

270.0 

3-22-20 

33 

595 

30  0 

270  0 

33 

941 

120.0 

270.0 

4-21-20 

34 

580 

30.0 

270.0 

34 

957 

120.0 

270.0 

5-21-20 

35 

591 

30  0 

270  0 

35 

966 

120.0 

270.0 

6-20-20 

36 

696 

30.0 

270.0 

36 

976 

120.0 

270  0 

7-20-20 

37 

613 

30.0 

270.0 

37 

995 

120.0 

270.0 

8-19-20 

38 

603 

30  0 

270.0 

38 

990 

120.0 

270.0 

9-18-20 

39 

619 

30.0 

270.0 

39 

1000 

120.0 

270.0 

Steer  575 

Steer 

574 

8-  5-17 

1 

217 

13.4 

75.8 

2.35 

1 

242 

18.0 

78.6 

240.0 

9-  4-17 

2 

227 

.2 

75.0 

236  5 

2 

244 

9.8 

90.0 

226.9 

10-  4-17 

3 

228 

109.0 

67.0 

3 

252 

23.4 

126.2 

62.5 

11-  3-17 

4 

237 

17.1 

135.0 

4 

264 

45.0 

141.0 

12-  3-17 

5 

246 

30  0 

143.9 

5 

268 

46.5 

150.0 

1-  2-18 

6 

257 

30.0 

164.5 

6 

284 

51.0 

164.5 

2-  1-18 

7 

275 

18.3 

165.0 

7 

309 

51.2 

165.0 

3-  3-18 

8 

283 

155.0 

8 

315 

11.5 

165.0 

4-  2-18 

9 

290 

150.0 

9 

315 

5.5 

165.0 

5-  2-18 

10 

308 

150.0 

10 

337 

174.0 

6-  1-18 

11 

304 

150.0 

11 

337 

174.0 

7-  1-18 

12 

307 

150.0 

12 

332 

180.0 

7-31-18 

13 

294 

150.0 

13 

322 

180.0 

8-30-18 

14 

298 

161.5 

14 

324 

193.0 

9-29-18 

15 

299 

165.0 

15 

321 

195.0 

10-29-18 

16 

294 

27.5 

152.5 

16 

316 

210.5 

1 1-28-18 

17 

295 

53.5 

151.0 

17 

309 

32.5 

192.5 

12-28-18 

18 

308 

59.0 

148.0 

18 

321 

60.0 

175.5 

1-27-19 

19 

328 

60.0 

150.0 

19 

333 

60.0 

180.0 

2-26-19 

20 

343 

60.0 

150.0 

20 

348 

60.0 

180.0 

3-28-19 

21 

358 

60.0 

150.0 

21 

368 

60.0 

180.0 

4-27-19 

22 

380 

60.0 

150.0 

22 

382 

60.0 

180.0 

5-27-19 

23 

394 

60.0 

154.0 

23 

412 

60.0 

180.0 

6-26-19 

24 

409 

60.0 

179.0 

24 

417 

60.0 

180.0 

7-26-19 

25 

426 

60.0 

180.0 

25 

445 

60.0 

180.0 

8-25-19 

26 

425 

60.0 

180.0 

26 

445 

60.0 

180.0 

9-24-19 

27 

436 

60.0 

180.0 

27 

432 

60.0 

180.0 

10-24-19 

28 

441 

60.0 

180.0 

28 

439 

60.0 

180.0 

11-23-19 

29 

447 

60.0 

180.0 

29 

435 

60.0 

180.0 

12-23-19 

30 

458 

60.0 

180.5 

30 

448 

60.0 

180.5 

1-22-20 

31 

472 

60.0 

180.0 

31 

453 

60.0 

180.0 

2-21-20 

32 

480 

60.0 

180.0 

32 

464 

60.0 

180.0 

3-22-20 

33 

496 

60.0 

180.0 

33 

474 

60.0 

180.0 

4-21-20 

34 

504 

60.0 

180.0 

34 

481 

60.0 

180.0 

5-21-20 

35 

515 

60.0 

180.0 

35 

499 

60.0 

180.0 

6-20-20 

36 

528 

60.0 

180.0 

36 

511 

60.0 

180.0 

7-20-20 

37 

538 

60.0 

180.0 

37 

518 

60.0 

180.0 

8-19-20 

38 

530 

60.0 

180.0 

38 

506 

60.0 

180.0 

9-18-20 

39 

537 

60.0 

180.0 

39 

518 

60.0 

180.0 

"Average  weight  of  last  ten  days  of  period. 


28  Missouri  Agricultural  Experiment  Station  Bulletin  51 


Table  13  (Continued). — Weight  in  Pounds  of  Animals  and  of  Feed  Consumed 
by  Thirty  Day  Periods 


Date 

beginning 
of  period 

Period 

No. 

Live 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Period 

No. 

Live 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Steer  573 

Steer 

572 

8-  5-17 

1 

197 

14.1 

64.9 

214.0 

1 

210 

16.4 

72.6 

215.4 

9-  4-17 

2 

220 

34.1 

102.3 

253.0 

2 

216 

22.1 

92.1 

42.3 

10-  4-17 

3 

237 

75.7 

108.4 

50.5 

3 

231 

28.8 

124.9 

11-  3-17 

4 

269 

105.8 

128.1 

4 

243 

30.0 

134.5 

12-  3-17 

5 

290 

87.5 

145.4 

5 

252 

30.0 

146.5 

1-  2-18 

6 

312 

72.0 

164.3 

6 

263 

30.0 

150.0 

2-  1-18 

7 

335 

62.5 

165  0 

7 

280 

16.3 

154.5 

3-  3-18 

8 

345 

30.0 

165  0 

8 

289 

155.0 

4-  2-18 

9 

354 

29.5 

165  6 

9 

295 

0.5 

150.0 

5-  2-18 

10 

366 

15.0 

192  0 

10 

305 

150.0 

6-  1-18 

11 

364 

6.0 

202.5 

11 

307 

150.0 

7-  1-18 

12 

369 

210.0 

12 

311 

150.0 

7-31-18 

13 

355 

210  0 

13 

306 

150.0 

8-30-18 

14 

367 

223  0 

14 

304 

165.0 

9-29-18 

15 

364 

225  0 

15 

309 

165.0 

10-29-18 

16 

361 

213  5 

16 

309 

179.0 

11-28-18 

17 

339 

9.5 

167  5 

17 

304 

182.0 

12-28-18 

18 

354 

30  0 

180  5 

18 

309 

180.0 

1-27-19 

19 

383 

30.0 

221.5 

19 

317 

187.0 

2-26-19 

20 

399 

30.0 

221.5 

20 

328 

210.0 

3-28-19 

21 

406 

43.0 

213.5 

21 

341 

210.0 

4-27-19 

22 

431 

60.0 

210.0 

22 

358 

210.0 

5-27-19 

23 

438 

60.0 

212.5 

23 

382 

206.0 

6-26-19 

24 

454 

60.0 

217.5 

24 

370 

210.5 

7-26-19 

25 

477 

60.0 

209.0 

25 

375 

210.0 

8-25-19 

26 

492 

60.0 

210.0 

26 

380 

210.0 

9-24-19 

27 

485 

60.0 

206.0 

27 

378 

210.0 

10-24-19 

28 

499 

60.0 

200.5 

28 

375 

210.0 

11-23-19 

29 

503 

60.0 

210.5 

29 

372 

224.5 

12-23-19 

30 

522 

60.0 

209.5 

30 

392 

241.5 

1-22-20 

31 

532 

60.0 

210.0 

31 

414 

254.0 

2-21-20 

32 

537 

60.0 

210.0 

32 

434 

270.0 

3-22-20 

33 

552 

60.0 

210.0 

33 

448 

270.0 

4-21-20 

34 

557 

60.0 

210.0 

34 

433 

238.5 

5-21-20 

35 

569 

60.0 

210.0 

35 

460 

270.0 

6-20-20 

36 

581 

60.0 

210.0 

36 

469 

270.0 

7-20-20 

37 

595 

60.0 

210.0 

37 

479 

270.0 

8-19-20 

38 

588 

60.0 

210.0 

38 

485 

270.0 

9-18-20 

39 

594 

60.0 

209.5 

39 

488 

270.0 

Steer  571 

8-  5-17 

1 

173 

24.5 

79.3 

82.8 

9-  4-17 

2 

181 

32.5 

103.0 

10-  4-17 

3 

200 

81.1 

96.9 

11-  3-17 

4 

237 

109.6 

111.0 

12-  3-17 

5 

264 

105.9 

123.3 

1-  2-18 

6 

288 

102.0 

134.8 

2-  1-18 

7 

321 

102.0 

136.5 

3-  3-18 

8 

355 

102.0 

135.0 

4-  2-18 

9 

383 

103.0 

142.0 

5-  2-18 

10 

412 

76.0 

177.5 

6-  1-18 

11 

433 

70.0 

189.0 

The  Influence  of  the  Plane  of  Nutrition 


29 


Table  13  (Continued). — Weight  in  Pounds  of  Animals  and  of  Feed  Consumed 
by  Thirty  Day  Periods 


Date 

beginning 
of  period 

Period 

No. 

Live 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Period 

No. 

Live 

weight 

Pounds 

Grain 

Pounds 

Hay 

Pounds 

Milk 

Pounds 

Steer  571 

7-  1-18 

(Cont.) 

12 

453 

75.0 

195.0 

7-31-18 

13 

469 

75.0 

195.0 

8-30-18 

14 

478 

75  0 

208.0 

9-29-18 

15 

483 

75.0 

210.0 

10-29-18 

16 

494 

88.5 

210.5 

11-28-18 

17 

499 

90.0 

163.0 

12-28-18 

18 

510 

90.0 

180.5 

1-27-19 

19 

536 

90.0 

207.0 

2-26-19 

20 

559 

90.0 

210.0 

3-28-19 

21 

502 

90.5 

210.0 

4-27-19 

22 

496 

90.0 

210.0 

5-27-19 

23 

612 

90.0 

210.0 

6-26-19 

24 

616 

88.5 

206.5 

7-26-19 

25 

635 

90.0 

210.0 

8-25-19 

26 

637 

89.9 

210.0 

9-24-19 

27 

617 

90.0 

213.0 

* 

10-24-19 

28 

627 

90.0 

230.0 

11-23-19 

29 

649 

90.0 

242.0 

12-23-19 

30 

667 

90.0 

240.0 

1-22-20 

31 

681 

88.5 

228.5 

2-21-20 

32 

699 

90.0 

240.0 

3-22-20 

33 

716 

90.0 

240.0 

4-21-20 

34 

727 

90.0 

240.0 

5-21-20 

35 

747 

90.0 

240.0 

6-20-20 

36 

755 

90.0 

240.0 

7-20-20 

37 

754 

90.0 

240.0 

8-19-20 

38 

752 

90.0 

240.0 

9-18-20 

39 

757 

90.0 

240.0 

30  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  14. — Dry  Matter  and  Organic  Matter  in  Feed  by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer  585 
1 

8.020 

7.655 

7.655 

4.464 

2 

4.714 

14.211 

4.548 

18.107 

13.559 

11.480 

3 

53.811 

48.683 

48.683 

28.392 

4 

84.158 

67.090 

76.090 

44 . 376 

5 

97.228 

87.906 

87.906 

51.267 

6 

26.974 

111.024 

100.449 

100.449 

58 . 582 

7 

127.800 

115.769 

115.769 

67.516 

8 

136.098 

123.498 

123.498 

72 . 024 

9 

136.098 

123.498 

123 . 498 

72 . 024 

10  

136.098 

123.498 

123.498 

72 . 024 

11 

136.098 

123.498 

123.498 

72.024 

12 

136.098 

123.498 

123.498 

72.024 

13 

136.098 

123.498 

123.498 

72 . 024 

14 

144.047 

131.017 

131.017 

76 . 409 

15  

10  616 

142.540 

10.231 

129.640 

139.871 

84 . 342 

16  

33.196 

151.424 

31.983 

138.174 

170.157 

102 . 605 

17  

61.760 

139.866 

60 . 208 

138.656 

188.864 

113.885 

18  

54 . 639 

143.105 

52.691 

129.375 

182.066 

115.430 

19 

54.330 

167.263 

52 . 394 

51.213 

103.617 

65.687 

20  

53.710 

168.654 

51.810 

152.515 

204 . 324 

129 . 542 

21 

53.724 

178.837 

51.847 

162.701 

214.548 

136.023 

22  

50 . 568 

164.763 

48.801 

148.733 

197.534 

125.236 

23  

30.431 

164.763 

29.311 

148.733 

178.044 

110.904 

24 

27.072 

165.125 

26.013 

149.135 

175.148 

109.100 

25 

27.072 

165.125 

26.013 

149.135 

175.148 

109.100 

26 

28.422 

164.641 

27.311 

157.199 

184.510 

114.931 

27 

40.609 

166.170 

39.021 

152.310 

191.331 

121.304 

28 

40 . 609 

166.170 

39.021 

152.310 

191.331 

121.304 

29 

38.357 

156.923 

36.857 

143.843 

180.700 

114.564 

30 

40.228 

172.617 

38.630 

158.217 

196.837 

124.795 

31  

39 . 846 

185.540 

38.217 

170.070 

208.287 

132.054 

32  

45.156 

193.915 

43.311 

176.576 

219.887 

139.408 

33  

66.404 

207 . 059 

63 . 690 

187.969 

251.659 

159.552 

34  

58 . 432 

201.112 

56.044 

182.562 

238.606 

151.276 

35  

48 . 702 

193.255 

46.712 

175.435 

222.147 

140.841 

36  

33 . 684 

193.255 

32.324 

175.435 

207.759 

129.413 

37 

26.769 

193.255 

25.721 

175.435 

201.156 

125.300 

38 

22 . 343 

193.255 

21.477 

175.435 

196.912 

118.738 

39  

13.406 

191.933 

12.886 

174.233 

187.119 

112.833 

40  

13.406 

202.915 

12.886 

184.205 

197.091 

118.846 

41  

33 . 063 

207 . 059 

31.781 

187.969 

219.750 

136.882 

42  

62.557 

210.216 

60.132 

191.418 

251.550 

159.483 

43  

59 . 770 

218.126 

57.487 

198.876 

256.363 

162.534 

44  

64 . 609 

221.997 

62.164 

201 . 753 

263.917 

167.323 

45 

67.736 

219.940 

65.172 

199.900 

265.072 

168.056 

46 

32.538 

219.940 

31.307 

199.900 

231.207 

144.019 

47  

44 . 566 

215.383 

42 . 880 

195.753 

238.633 

151.283 

48  

207 . 593 

187.163 

187.163 

109.143 

49 

206 . 844 

187.950 

187.950 

109.612 

50 

219.508 

199.066 

199.066 

116.095 

51  

222.130 

197.050 

197.050 

114.920 

52  

222.130 

197.050 

197.050 

114.920 

53  

234 . 484 

208.361 

208.361 

121.516 

The  Influence  of  the  Plane  of  Nutrition 


31 


Table  14  (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer~685  (Cont.) 

54  

236.436 

210.176 

210.176 

122.575 

55  

243.117 

215.919 

215.919 

125.924 

5fi  

244.898 

215.688 

215.688 

125.789 

57  

289.470 

254.970 

254.970 

148.698 

58  

. . 279 . 340 

246.020 

246.020 

143.479 

5Q  

290.748 

256.060 

256.060 

143.502 

fiO  

290.740 

256.060 

256.060 

149.354 

fil  

281.412 

249.172 

249.172 

145.317 

«2  

291.199 

260.959 

260.959 

152.191 

88  

297.227 

. .267. 117 

267.117 

155.783 

84 

319.520 

287.150 

287.150 

167.466 

319  520 

287.150 

287.150 

167.466 

66 

27.663 

280.876 

26.661 

252.426 

279.087 

168.298 

67 

52.877 

304.072 

50.961 

275.429 

326.390 

196.813 

68 

51.105 

275.950 

49.254 

254.840 

304 . 094 

192.796 

69 

53.460 

297.126 

51.281 

269.746 

321.027 

199.968 

70 

53.460 

320.353 

51.281 

290.853 

342.134 

213.115 

71 

53.460 

320.353 

51.281 

290.853 

342.134 

213.115 

72 

53.460 

320.353 

51.281 

290.853 

342.134 

213.115 

73 

54.787 

320.353 

51.513 

290.853 

342.366 

283.260 

74 

55.552 

319.243 

53.227 

289.813 

343.040 

213.680 

75 

55.552 

320.353 

53.227 

290.853 

344.080 

214.327 

76 

55.552 

324.764 

53.227 

294 . 509 

347.836 

216.667 

77 

55.552 

338 . 393 

53.227 

305.928 

358.155 

223.095 

78 

53.233 

338.393 

50.292 

305.928 

355.221 

221.267 

Steer  579 

1 

2.337 

2.230 

2.230 

1.301 

2 

6.835 

13.297 

6.595 

12.687 

19.289 

12.406 

3 

21.317 

41.92 

20.434 

37.937 

58.371 

37.556 

4 

26.663 

56.577 

25.563 

51.153 

76.716 

49.359 

5 

33.813 

69.177 

32.228 

62.544 

94 . 772 

60.976 

6 

39.738 

82.904 

38.192 

75.032 

113.224 

72.848 

7 

39.738 

66.513 

38.172 

57.667 

95.839 

61.662 

8 

39.738 

108.874 

38.192 

98 . 804 

136.996 

88.143 

9 

40.674 

108.874 

39.156 

98.804 

137.960 

88.763 

10 

40.614 

110.697 

39.152 

100.447 

139.599 

89.817 

11 

40.614 

108.874 

39.152 

98.804 

137.956 

88.761 

12 

40.614 

108.874 

39.152 

98.804 

137.956 

88.761 

13 

40.614 

108.874 

39.152 

98.804 

137.956 

88.761 

14 

45.162 

114.739 

43 . 540 

104.549 

148.189 

95.345 

15 

64.604 

132.805 

62.276 

120.795 

183.071 

117.788 

16 

68.975 

168.623 

66.455 

154.178 

220.633 

141.955 

17 

61.512 

167.275 

59.199 

153.855 

213.054 

137.079 

18 

54 . 639 

167.253 

52.691 

151.213 

203 . 904 

131.191 

19 

58.361 

171.440 

56.282 

155.000 

211.282 

133.953 

20 

68.488 

180.275 

66.066 

162.906 

228.972 

147.321 

21 

67.142 

179.823 

64 . 796 

162.344 

227.140 

146.142 

22 

67.142 

178.459 

61.796 

161.099 

225.895 

145.341 

23 

67.129 

178.459 

64.642 

161.099 

225.741 

145.242 

24 

07.677 

179.865 

65.031 

161.545 

226.576 

145.779 

25 

66.766 

177.97 

64.155 

160.730 

224.885 

144.691 

32  Missouri  Agricultural  Experiment  Station  Bulletin  51 


Table  14  (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer  579  (Cont.) 

26 

61.361 

165.772 

58.961 

151.134 

210.095 

135.175 

27 

67.677 

166.17 

65.031 

15^.310 

217.341 

139.837 

28 

67.677 

166.17 

65.031 

152.31 

217.341 

139.837 

29 

67.677 

166.17 

65.031 

152.31 

217.341 

139 . 837 

30 

73.553 

173.084 

70.868 

158.654 

229.523 

147.675 

31 

71.727 

174.565 

68.796 

159.945 

228.741 

147.171 

32 

79.687 

184.228 

76.431 

167.765 

244.196 

157.116 

33 

79.687 

198.794 

76.431 

180.464 

256.895 

165.286 

34 

82.341 

194.189 

78.976 

176.279 

255.255 

164.231 

35 

92.967 

207.059 

89.168 

187.969 

277.137 

178.309 

36 

84.667 

206.591 

81.239 

187.541 

268.770 

172.927 

37 

83 . 882 

211.187 

80.602 

191.717 

272.319 

175. 2’0 

38 

81.771 

206.591 

78.600 

187.541 

266.141 

171.235 

39 

67.068 

207.059 

64.469 

187.969 

252.438 

160.046 

40 

67.068 

207.059 

64.469 

187.969 

252.438 

160.046 

41 

76.865 

207.059 

73.885 

187.261 

261.854 

166.015 

42 

126.453 

215.797 

121.550 

1»6.499 

318.049 

204.633 

43 

121.745 

232.210 

117.109 

211.670 

328.779 

211.536 

44 

131.020 

232.557 

126.062 

211.336 

337.398 

217.082 

45 

146.614 

231.830 

141.067 

210.700 

351.767 

226.327 

46 

106.958 

233.635 

102.910 

212.335 

315.245 

202.829 

47 

59.722 

233.635 

57.462 

212.335 

269.797 

171.051 

48 

65.942 

235.717 

63.459 

212.527 

275.986 

174.975 

49 

20.256 

269.474 

19.495 

244.911 

264.406 

159.437 

50 

282.572 

256.351 

256.351 

149 . 504 

51 

291.554 

258.634 

258.634 

150.835 

52 

291.554 

258.634 

258.634 

150.835 

53 

23.476 

308.734 

22.572 

274.357 

296.929 

179.048 

54 

1.805 

314.352 

1.736 

279.442 

281.178 

163.983 

*»5  

233.442 

207.339 

207.339 

120.920 

56  

268.290 

236.290 

236.290 

137.805 

57 

290.760 

256 . 080 

256 . 080 

149  346 

58  

309 . 700 

272.760 

272.760 

159.074 

59 

278.747 

240.907 

240.907 

140.497 

60  

278.747 

240.907 

240.907 

140.497 

61  

316.923 

280.723 

280.723 

163.718 

62  

256.276 

229.544 

229 . 544 

133.870 

63  

289.234 

259.924 

259.924 

151.588 

64  

318.530 

286.250 

285.250 

166.941 

65  

319.620 

287.150 

287.150 

167.466 

66 

7.489 

19.520 

7.218 

287.150 

294.368 

171.675 

67 

51.991 

320.059 

50.107 

289.896 

340.003 

198.290 

68 

53.442 

317.243 

51.279 

287.753 

339.032 

211.183 

69 

53.460 

293.057 

51.281 

266.047 

317.328 

197.664 

70 

53.460 

293.057 

51.281 

266.047 

317.328 

197.664 

71 

53.460 

293.057 

51.281 

266.047 

317.328 

197.664 

72 

53.460 

293.057 

51.281 

266.047 

317.328 

197.664 

73 

54.556 

317.243 

52.576 

287.753 

340.329 

205.218 

74 

54.629 

291.776 

52.340 

264.906 

317.246 

191.299 

75 

55.555 

325.603 

53.227 

295.275 

348.502 

210.147 

76 

55.555 

338.393 

53.227 

305.928 

359.155 

216.570 

77 

55.555 

338.393 

53.227 

305.928 

359.155 

216.570 

78 

53.233 

338.393 

50.293 

305.928 

356.221 

214.801 

The  Influence  of  the  Plane  of  Nutrition 


33 


Table  14  (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer  528 
1 

2.292 

2.187 

2.187 

1.275 

2 

13.199 

12.836 

12.735 

12.247 

24.982 

17.377 

3 

24.449 

48.210 

18.829 

43.620 

62.449 

43.440 

4 

61.320 

70.133 

58.791 

63.410 

122.201 

85.003 

5 

97.749 

94.913 

93.861 

85.814 

179.675 

124.982 

6 

105.963 

110.548 

101.839 

100.052 

201.891 

140.435 

7 

92.871 

109.557 

89.255 

99.249 

188.504 

131.123 

8 

106.422 

108.884 

102.297 

98.804 

201.101 

139.886 

9 

108.485 

108.485 

104.436 

104.436 

208.872 

145.291 

10 

108.311 

107.972 

104.411 

97.977 

202.388 

140.781 

11 

108.311 

108.884 

104.411 

98.804 

203.215 

141.356 

12 

108.311 

108.884 

104.411 

98.804 

203.215 

141.356 

13 

108.311 

108.884 

104.411 

98.804 

203.215 

141.356 

14 

108.322 

120.711 

104.430 

109.791 

214.221 

149.012 

15 

108.374 

138.046 

104.467 

125.436 

229.903 

159.921 

16 

131.778 

141.910 

126.964 

130.344 

257.308 

178.983 

17 

135.684 

167.303 

130.728 

153.883 

284.611 

197.975 

18 

136.606 

167.253 

131.736 

151.213 

282.949 

196.819 

19 

- 140.308 

171.440 

135.309 

155.000 

290.309 

210.939 

20 

147.715 

181.653 

142.492 

164.154 

306.646 

213.303 

21 

147.658 

179.823 

142.496 

162.244 

304 . 740 

211.977 

22 

147.658 

178.459 

142.496 

161.099 

303.595 

211.181 

23 

147.668 

178.459 

142.194 

161.099 

303 . 293 

210.971 

24 

148.916 

178.865 

143.094 

161.545 

304 . 639 

211.907 

25 

148.916 

178.865 

143.094 

161.545 

304.639 

211.907 

26 

148.916 

179.161 

143.094 

163.418 

306.512 

213.210 

27 

148.916 

166.170 

143.094 

152.310 

295.404 

205.483 

28 

148.916 

166.638 

143.094 

152.738 

295.832 

205.781 

29 

162.008 

174.495 

155.674 

159.945 

315.619 

219.545 

30 

160.901 

186.971 

154.469 

171.381 

325.840 

226.654 

31 

159.286 

294 . 778 

152.773 

178.528 

331.301 

230.459 

32 

159.286 

197.634 

152.773 

179.975 

332.748 

231.459 

33 

159.286 

207.974 

152.773 

188.794 

341.567 

237.594 

34 

159.286 

207.059 

152.773 

187.969 

340.742 

237.020 

35 

159.286 

207.547 

152.773 

188.407 

340.180 

236.629 

36 

159.615 

207.059 

153.136 

187.969 

341.105 

237.273 

37 

169.556 

210.292 

162.924 

190.892 

353.816 

246.114 

38 

160.889 

207 . 059 

154.652 

187.969 

342.621 

238 . 327 

39 

100.889 

207.059 

154.652 

187.969 

342.621 

238.327 

40 

168.125 

208.878 

161.610 

189.218 

351.228 

244.314 

41 

175.158 

219.033 

168.366 

198.833 

367.199 

255.424 

42 

186.821 

215.766 

179.570 

196.467 

376.037 

261.571 

43 

200.193 

225.637 

192.791 

205.717 

398.508 

277.202 

44 

200.504 

221.956 

192.914 

201.706 

394 . 620 

274.498 

45 

200.504 

219.940 

192.914 

199.900 

392.814 

273.241 

46 

200.504 

219.940 

192.914 

199.900 

392.814 

273 . 24 1 

47 

200 . 504 

219.940 

192.914 

199.900 

392.814 

273.241 

48 

201.409 

221.858 

193.821 

200.028 

393.849 

273.961 

49 

132.998 

270.41 6 

129.924 

145.763 

375.687 

241.717 

50 

87.287 

290.807 

84 . 007 

263.791 

347.798 

223 . 773 

51 

25.168 

305 .412 

24.211 

270.932 

295. 143 

177.971 

52 

27.026 

305.412 

26 . 539 

270.932 

297.471 

179.375 

34  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  14  (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer  528  (Cont.) 

53  

27.085 

306.048 

26.042 

271.986 

298.028 

179.711 

182.640 

54 

27.085 

311.443 

26 . 042 

276.843 

302.885 

55 

78.548 

307.178 

75 . 522 

272.820 

348.342 

220.848 

56 

107.548 

311.360 

103.381 

274.210 

377.591 

239.393 

57 

105.910 

314.960 

101.791 

277.380 

378.171 

240.394 

58 

105.576 

317.150 

101.468 

279.310 

380.778 

241 .413 

59 

132.253 

317.150 

127.092 

279.310 

406.392 

261.472 

60 

131.819 

317.150 

126 . 665 

297.310 

405.975 

261.204 

61  

131 .006 

319.497 

126.089 

283.017 

409.106 

263.219 

266.241 

62  

130.710 

321.207 

125.894 

287.909 

413.803 

63  

131.769 

316.860 

126.888 

284.760 

411.648 

264 . 854 

64  

132.111 

319.520 

127.268 

287.150 

414.418 

266.637 

65 

132.181 

319.520 

127.392 

287.150 

414.542 

266.716 

66  

132.181 

319.520 

127.392 

287.150 

414.542 

266.716 

67 

132.181 

320.062 

127.392 

289.899 

417.291 

268.485 

68 

132.249 

320.273 

126.891 

290.783 

417.670 

268.731 

69 

133.678 

320.273 

128.232 

290.783 

419.015 

269.594 

70 

133.678 

320.273 

128.232 

290.783 

419.015 

269 . 594 

71 

133.678 

320.273 

128.232 

290.783 

419.015 

269.594 

72 

122 . 678 

320.273 

128.232 

290.783 

419.015 

269 . 594 

73 

136.974 

320.027 

131.291 

290.783 

422.074 

271.562 

74 

136.573 

320.027 

130.850 

290.783 

421.633 

271.279 

75 

138.888 

320.027 

130.850 

290 . 783 

421.633 

271.279 

76 

138.888 

320.061 

130.850 

290.246 

421.096 

270.933 

77 

136.110 

327.113 

130.407 

295.730 

426.137 

274.177 

78 

133.085 

338.093 

125.734 

305.928 

431.662 

277.731 

Steer  578 
1 

22.361 

104.971 

21.945 

96.528 

118.023 

74 . 827 

2 

39 . 593 

102.156 

38 . 058 

92.737 

130.795 

82 . 924 

3 

64.522 

124.266 

62.020 

111.347 

173.367 

111.544 

4 

80.281 

131.978 

77.222 

120.328 

197.550 

127.104 

5 

67.831 

180.588 

65.270 

164.178 

229.448 

147.627 

6 

72.191 

150.751 

69 . 459 

137.011 

206.470 

132.843 

7 

112.607 

151.207 

109.855 

137.427 

247.232 

159.101 

8 

25.852 

151.207 

24 . 874 

137.427 

162.301 

102.899 

9 

19.304 

152.693 

18.577 

137.668 

156.245 

94.216 

10 

8.998 

176.927 

8.660 

160.696 

169.356 

98.768 

11 

180.748 

161.434 

161.434 

94.148 

12 

180.485 

160.100 

160.100 

93.370 

13 

180.485 

160.100 

160.100 

93.370 

14 

192.763 

171.302 

171.302 

99 . 903 

15 

194.679 

173.056 

173.056 

100.926 

16 

1.805 

159.232 

1.736 

144.700 

146.436 

85.400 

17  

204 . 744 

180.211 

180.211 

105.099 

18 

199.535 

175.740 

175.740 

102.492 

19 

211.430 

186.214 

186.214 

108 . 600 

20 

211.430 

186.214 

186.214 

108.600 

21 

211.430 

186.214 

186.214 

108 . 600 

22  

212.996 

188.682 

188.682 

110.039 

23 

215.343 

193.013 

193.013 

112.565 

24 

217.377 

195.347 

195.347 

113.926 

The  Influence  of  the  Plane  of  Nutrition 


35 


TableT 4 (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer]578  (Cont.) 

25 

14.981 

238.663 

14.438 

214.483 

228.921 

133.507 

26 

26.438 

239.560 

25.480 

215.280 

240.760 

145. 178 

27 

26.879 

239.560 

25.905 

215.280 

241.185 

145.435 

28 

26.438 

216.989 

25.480 

194.367 

219.847 

132.568 

29 

26.718 

240.187 

25.633 

218.066 

243.699 

146.950 

30 

26.727 

240.187 

25.638 

218.066 

243.704 

146.954 

31 

26.727 

240.187 

25.638 

218.066 

243 . 704 

146.954 

32 

26.727 

240.187 

25.638 

218.066 

243 . 704 

146.954 

33 

26.727 

240.187 

25.638 

218.066 

243 . 704 

146.954 

34 

27.065 

240.187 

26.652 

218.006 

244.718 

147.565 

35 

27.778 

240.187 

26.614 

218.066 

244.680 

147.542 

36 

27.778 

240.187 

26.614 

218.066 

244 . 680 

147.542 

37 

27.778 

244 . 278 

26.614 

221.518 

248.132 

149.624 

38 

27.778 

253.795 

26.614 

229.446 

256.060 

154.404 

39 

26.616 

253.795 

25.147 

229.446 

254.593 

153.520 

Steer  577 

1 

18.999 

73.079 

18.185 

66.338 

84.523 

53.588 

2 

29.488 

94.806 

28.344 

85.969 

114.313 

72.474 

3 

63.460 

110.490 

61.000 

100.619 

161.619 

108.204 

4 

84.664 

121.482 

81.436 

120.317 

201.750 

135.072 

5 

93.583 

180.626 

90.041 

163.214 

253 . 255 

169.554 

6 

93 . 583 

150.736 

90.041 

136.996 

237.037 

158.696 

7 

95.834 

151.206 

92.208 

137.426 

229 . 634 

153.740 

8 

106.958 

151.206 

102.910 

137.426 

240.336 

160.905 

9 

107.214 

152.228 

103.176 

137.248 

240.424 

160.964 

10 

82 . 340 

179.679 

79.245 

163.279 

242.524 

156.040 

11 

80.991 

181.545 

77.947 

164.658 

242.605 

156.092 

12 

80.889 

194.372 

77.812 

172.419 

250.231 

160.999 

81.809 

194.372 

78.276 

172.419 

250.695 

161.297 

14 

81.260 

206.674 

78.130 

183.643 

261.773 

168.425 

15 

81.260 

208.596 

78.130 

185.426 

263.556 

169.572 

16 

102.026 

224.104 

98.096 

202 . 282 

300.378 

193.263 

17 

107.569 

185.009 

103.402 

162.939 

266.341 

171.364 

18 

105.926 

185.009 

101.801 

162.939 

264 . 740 

170.334 

19 

105.602 

203.512 

101.493 

179.233 

280.726 

180.619 

20 

105.460 

211.451 

101.337 

186.221 

287.558 

185.015 

21 

105.460 

211.451 

101.337 

186.221 

287.558 

185.015 

22 

103.973 

202.745 

100.071 

179.586 

279.657 

179.931 

23 

104.567 

211.007 

100.714 

191.817 

292.531 

188.214 

24 

106.204 

212.964 

102.268 

191.384 

293 . 652 

188.936 

25 

101 . 308 

212.964 

97.469 

191.384 

288 . 853 

185.848 

26 

105.765 

212.174 

101.933 

190.584 

292.517 

188.205 

27 

94.751 

200.989 

91.317 

180.629 

271.946 

174.970 

28 

105.765 

229.396 

101.933 

207.780 

309.713 

199.269 

29 

106.849 

240.173 

102.528 

218.043 

320.571 

206 . 255 

30 

106.917 

240.173 

102.560 

218.043 

320.603 

206 . 276 

31 

106.917 

240. 173 

102.560 

218.043 

320.603 

206 . 276 

32 

106.917 

240.173 

102.560 

218.043 

320.603 

206.276 

33 

106.917 

240.173 

102.660 

218.043 

320.603 

206 . 276 

34 

110.112 

240.173 

105.556 

218.043 

323 . 599 

208.204 

35 

111.110 

240.173 

106.454 

218.043 

324.497 

208.781 

36  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  14  (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer  577  (Cont.) 

36 

111.110 

240.173 

108.454 

218.043 

324.497 

208.781 

37 

111.110 

244.278 

106.454 

221.518 

327.972 

211.017 

38 

111.110 

253.795 

106.454 

229.445 

335.900 

216.118 

39 

106.465 

253.795 

100.597 

229.464 

330.051 

212.355 

Steer  575 

1 

12.158 

69.762 

11.512 

63.329 

74.841 

46.618 

2 

.182 

69.026 

.175 

62.661 

62.836 

36.646 

3 

115.261 

106.173 

106.173 

61.920 

4 

15.236 

151.560 

14.659 

140.360 

155.019 

90.407 

5 

26.742 

142.210 

26.730 

129.225 

154.955 

90.370 

6 

26.742 

150.379 

26.730 

136.669 

162.399 

94.711 

7 

16.314 

151.206 

15.696 

137.426 

153.122 

89 . 008 

R 

142.028 

129.078 

129.078 

75.278 

g 

147.389 

133.779 

133.779 

78 . 020 

10 

139.213 

125.591 

125.591 

73 . 245 

11 

137.162 

124.453 

124.453 

72.581 

12 

138.828 

123.158 

123  158 

71.826 

13 

138.828 

123.158 

123.158 

71.826 

14 

149.691 

133.024 

133 . 024 

77.580 

15 

152.834 

136.004 

136.044 

79.318 

16 

24.831 

140.477 

23 . 875 

124.735 

148.610 

94.219 

17 

47.748 

178.010 

45.892 

162.140 

208.032 

131.892 

18 

52.087 

130.398 

50.059 

114.848 

164.907 

104.551 

19 

52.798 

136.154 

50.744 

120.404 

171.148 

108.508 

20 

52 . 728 

136.154 

50.668 

120.404 

171.072 

108.460 

21 

52 . 728 

136.154 

50.668 

120.404 

171.072 

108.460 

22 

52.419 

133.109 

50.452 

117.913 

168.365 

106.743 

23 

52.284 

137.308 

50.357 

123.075 

173.432 

109.956 

24 

53 . 047 

158.877 

51.084 

142.777 

193.861 

122.908 

25 

52.848 

159.736 

50.911 

143.556 

194.467 

123.292 

26 

52.877 

159.736 

50.961 

143.556 

194.517 

123.324 

27 

52.877 

159.736 

50.961 

143.556 

194.517 

123.324 

28 

52.887 

149.840 

50.961 

144.933 

195.894 

124.197 

29 

21.274 

160.112 

19.105 

145.362 

164.467 

104.272 

30 

53.460 

160.552 

51.281 

145.762 

197.043 

124.925 

31 

53.460 

160.552 

51.281 

145.762 

197.043 

124.925 

32 

53.460 

160.552 

51.281 

145.762 

197.043 

124.925 

33 

53.460 

160.552 

51.281 

145.762 

197.043 

124.925 

34 

54.556 

160.552 

52.578 

145.762 

198.340 

125  748 

35 

55.555 

160.552 

53.227 

145.762 

198.989 

126.159 

36 

55.555 

160.552 

53.227 

145.762 

198.989 

126.159 

37 

55.555 

162.844 

53.227 

147.764 

200.991 

127.428 

38 

55.555 

169.196 

53.227 

152.964 

206.191 

130.725 

39 

53.233 

169.196 

50.293 

152.964 

203.257 

128.845 

Steer  574 

1 

16.086 

72.337 

15.462 

65.667 

81.129 

51.436 

2 

8.758 

82.829 

8.419 

75.192 

83.611 

50.417 

3 

20.911 

118.144 

20.100 

107.764 

127.864 

81.066 

4 

54.842 

132.642 

53.328 

120.860 

174.188 

110.435 

5 

41.451 

1 

139.717 

39 . 882 

127.064 

166.946 

105.844 

The  Influence  of  the  Plane  of  Nutrition 


37 


Table  14  (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer  574  (Cont.) 

6 

45.109 

150.755 

43 . 389 

137.013 

180.402 

114.375 

7 

45.686 

151.215 

43.959 

137.431 

181.390 

115.001 

8 

10.198 

151 .215 

9.810 

137.431 

147.241 

88.786 

9 

4.942 

152.268 

4.762 

137.288 

142.050 

82.844 

10 

160.321 

145.659 

145.659 

84 . 948 

11 

159.116 

144.338 

144.338 

84.178 

12 

166  605 

147.795 

147.795 

86.194 

13 

166  605 

147.795 

147.795 

86.194 

14 

169  588 

150.701 

150.701 

87.889 

15 

180.783 

160.703 

160.703 

93 . 722 

16 

262.436 

240.717 

240.717 

140.386 

17 

29.015 

169.589 

27.887 

149.359 

177.246 

110.407 

18 

52.969 

154.553 

50.906 

136.166 

187.072 

118.604 

19 

52.799 

158.582 

50.736 

139.662 

190.398 

120.712 

20 

52.730 

158.582 

50.668 

139.662 

190.330 

120.669 

21 

52.730 

158.582 

50.668 

139.662 

190.330 

120.669 

22 

52.823 

162.733 

50.856 

144.498 

195.354 

123.954 

23 

52.284 

160.518 

50.357 

143.869 

194.226 

123.139 

24 

53.047 

159.736 

51.081 

143.556 

194.637 

123.400 

25 

52.850 

159.736 

50.913 

143.556 

194.469 

123.293 

26 

52.877 

159.736 

50.961 

143.556 

194.517 

123.324 

27 

52.877 

159.736 

50.961 

143.556 

194.517 

123.324 

28 

52.877 

185.616 

50.961 

170.530 

221.491 

140.425 

29 

53.402 

160.127 

51 .250 

145.377 

196.627 

124.662 

30 

53 . 460 

160.571 

51.281 

145.781 

197.062 

124.937 

31 

53.460 

160.571 

51.281 

145.781 

197.062 

124.937 

32 

53.460 

160.571 

51.281 

145.781 

197.062 

124.937 

33 

53.460 

160.571 

51.281 

145.781 

197.062 

124.937 

34 

54.556 

160.571 

52.578 

145.781 

198.359 

125.760 

35 

55.556 

160.571 

52.578 

145.781 

199.008 

126.171 

36 

55.555 

160.571 

53.227 

145.781 

199.008 

126.171 

37 

55.555 

162.844 

53.227 

147.764 

200.991 

127.428 

38 

55.555 

169.196 

53.227 

152.964 

206.191 

130.725 

39 

53.233 

169.196 

50.293 

152.964 

203.257 

128.865 

Steer  573 

1 ' 

12.605 

59.714 

12.117 

54 . 205 

66.322 

42.048 

2 

34.491 

94.115 

29 . 300 

85.441 

114.741 

72.746 

3 

67.653 

101.288 

65.030 

92.233 

157.263 

105.288 

4 

94.395 

120.418 

90.791 

109.788 

200.579 

139.523 

5 

77.992 

134.305 

75.042 

122.030 

197.072 

131.940 

6 

65.162 

138.195 

62.733 

130.822 

199.555 

128.394 

7 

55.708 

151.216 

53 . 600 

137.436 

191.036 

123.913 

8 

26.740 

151.216 

25.728 

137.436 

163.164 

103.446 

9 

26.458 

152.228 

25.462 

137.248 

162.710 

103.158 

10 

13.499 

193.531 

12.992 

177.521 

190.513 

114.879 

11 

5.399 

185.064 

5.196 

168.046 

173.242 

101.035 

12 

194.340 

172.390 

172.390 

100.538 

13 

194.340 

172.390 

172.390 

100.538 

14 

206  712 

183 . 652 

183 . 652 

107.106 

15 

208.516 

185.426 

185.426 

108.140 

16 

196.976 

174.949 

174.949 

102.030 



38  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  14  (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounds 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer  573  (Cont.) 

17 

8.437 

147.573 

8.108 

129.973 

137.081 

82.660 

18 

26.485 

159.023 

25.453 

140.053 

165.506 

103.094 

19 

26.401 

195.150 

25.374 

171.870 

197.244 

122.863 

20 

26.365 

195.150 

25.334 

171.870 

197.204 

122.839 

21 

36.957 

187.201 

35.479 

164.871 

200.350 

127.022 

22 

52.070 

186.347 

50.103 

165.079 

215.182 

136.425 

23 

52.285 

189.491 

50.358 

169.844 

220.202 

139.608 

24 

53 . 046 

193.005 

51.081 

173.465 

224.536 

142.356 

25 

52.849 

185.425 

50.912 

166.635 

217.547 

137.925 

26 

52.877 

186.335 

50.961 

167.455 

218.416 

138.476 

27 

52.877 

182.819 

50.961 

164.299 

215.260 

136.475 

28 

52 . 877 

178.251 

50.961 

161.515 

212.476 

134.710 

29 

53.447 

186.960 

51.277 

169.782 

221.019 

140.126 

30 

53 . 460 

170.905 

51.281 

169.188 

220.469 

139.777 

31 

53.460 

171.309 

51.281 

169.588 

220.869 

140.031 

32 

53.460 

171.309 

51.281 

169.588 

220.869 

140.031 

33 

53.460 

171.309 

51.281 

169.588 

220.869 

140.031 

34 

54.556 

171.309 

52.578 

169.588 

222.166 

140.853 

35 

55.555 

171.309 

53.227 

169.588 

222.815 

141.265 

36 

55.555 

171.309 

53.227 

169.588 

222.815 

141.265 

37 

55.555 

189.994 

53.227 

172.293 

225.520 

142.980 

38 

55.555 

197.396 

53.227 

178.458 

231.785 

146.952 

39 

53.233 

197.396 

50.293 

178.458 

228.751 

145.028 

Steer  572 

1 

14.652 

66.795 

14 . 084 

60.635 

74.719 

47.372 

2 

19.763 

48.768 

18.997 

76.952 

95.949 

60.832 

3 

25.738 

116.754 

24.740 

106.329 

131.069 

83.098 

4 

26.760 

116.404 

25.741 

115.288 

141.029 

89.412 

5 

26.731 

135.427 

25.729 

123.064 

148.793 

94.335 

6 

26.741 

164.213 

25.729  % 

151.683 

177.412 

112.479 

7 

14.535 

141.583 

13.985  ' 

128.630 

142.665 

86.027 

8 

103.436 

90.486 

90.486 

52.771 

9 

.445 

138.389 

.428 

124.779 

125.207 

73.020 

10 

138.416 

125.794 

125.794 

73.363 

11 

137.202 

124.236 

124.236 

72 . 454 

12 

138.838 

123.158 

123.158 

71.826 

13 

138.838 

123.158 

123.158 

71.826 

14 

152.948 

135.909 

135.909 

79.262 

15 

152.973 

135.983 

135.983 

79.305 

16 

165.518 

147.204 

147.204 

85 . 849 

17 

165.933 

146.688 

146.688 

85.546 

18 

163.897 

145.077 

145.077 

84 . 609 

19 

170.337 

150.687 

150.687 

87.880 

20 

191 .319 

169.249 

169.249 

98.706 

21 

191.319 

169.249 

169.249 

98.706 

22 

188.313 

167.389 

167.389 

97.621 

23 

189.891 

164.650 

164.650 

96.024 

24 

186.759 

167.839 

167.839 

97.884 

25 

186.314 

167.455 

167.455 

97.660 

26 

186.333 

167.455 

167.455 

97.660 

27 

186.333 

167.455 

167.455 

97.660 

The  Influence  of  the  Plane  of  Nutrition 


39 


Table  14  (Continued). — Dry  Matter  and  Organic  Matter  in  Feed 
by  30-Day  Periods 


Period 

Dry 
matter 
in  grain 
Pounsd 

Dry 
matter 
in  hay 
Pounds 

Organic 
matter 
in  grain 
Pounds 

Organic 
matter 
in  hay 
Pounds 

Total 

organic 

matter 

Pounds 

Digestible 

organic 

matter 

Pounds 

Steer  572  (Cont.) 

28 

177.007 

161.031 

161.031 

93.913 

29 

199.681 

181.281 

181.281 

105.723 

30 

214.805 

195.015 

195.015 

113.733 

31 

225.909 

205.099 

205 . 099 

119.614 

32 

240  613 

218.433 

218.433 

127.390 

33 

240  613 

218.433 

218.433 

127.399 

34 

212.135 

192.595 

192.595 

112.321 

35 

240  613 

218.433 

218.433 

127.390 

36 

240  613 

218.433 

218.433 

127.390 

37.  . . 

244  276 

221.518 

221.518 

129.199 

38.  . . 

253  795 

229.446 

229.446 

133.813 

39 

253.795 

229.446 

229.446 

133.813 

Steer  571 

1 

21.901 

72.984 

21.025 

56.247 

87.272 

55 . 292 

2 

29.045 

94.772 

27.918 

86 . 032 

113.950 

72.244 

3 

72.497 

96.069 

69 . 688 

87.970 

157.658 

109.666 

4 

97.728 

104.381 

94 . 004 

95.171 

139.175 

96.801 

5 

94 . 529 

127.521 

90.957 

117.834 

208.791 

145.235 

6 

90.913 

123.562 

87.472 

112.292 

199.764 

138.955 

7 

90.913 

125.082 

87.472 

113.682 

201.154 

139.922 

8 

90.913 

127.725 

87.472 

112.445 

199.917 

139.062 

9 

127.496 

95.030 

122.696 

85.680 

108.376 

144.946 

10 

68.385 

163.855 

65.815 

148.935 

214.750 

136.152 

11 

62.991 

172.918 

60.624 

156.704 

217.328 

137.786 

12 

67.402 

180.741 

64 . 838 

160.371 

225.209 

142.783 

13 

86.448 

180.741 

83.134 

160.371 

243.505 

154.382 

14 

67.705 

193.738 

65.097 

172.265 

237.362 

150.488 

15 

67.705 

194.670 

65.097 

173.040 

238.157 

150.979 

16 

79 . 906 

193.852 

76 . 829 

172.117 

248.946 

160.171 

17 

80.680 

143.604 

77.554 

126.474 

204 . 028 

131.271 

18 

79 . 688 

159.023 

76.581 

140.053 

216.634 

139.382 

19 

79 . 204 

182.380 

76.122 

160.620 

236.742 

152.319 

20 

79.095 

185.009 

76.003 

162.939 

238.942 

153.735 

21 

78.346 

185.009 

75.236 

162.939 

238.175 

153.241 

22 

78.263 

183.664 

67.282 

165.090 

232.372 

149.508 

23 

78.425 

187.272 

75.535 

167.851 

243.386 

156.595 

24 

78.236 

183.263 

75.394 

164.693 

240.087 

154.472 

79.228 

186.335 

76.287 

167.455 

243.742 

156.824 

26 

79.235 

186.335 

76.364 

167.455 

243.819 

156.873 

27 

79.316 

189.013 

76.442 

169.863 

246.305 

158.473 

28 

79.316 

204 . 484 

76.442 

1S5.209 

261.651 

168.346 

29 

80. 160 

215.246 

76.905 

195.416 

272.321 

172.651 

30 

80.188 

213.455 

76.920 

193.795 

270.615 

171.633 

31 

78.852 

208.103 

75.639 

184.533 

260.172 

164.949 

32 

80.188 

208.103 

76.920 

184.533 

261.453 

165.761 

33 

80.188 

208.103 

76.920 

184.544 

261.464 

165.768 

34 

82.282 

208. 103 

78.865 

184.544 

263 . 409 

167.001 

35 

83.333 

208. 103 

79.841 

184.544 

264.385 

167.620 

36 

83.333 

208.103 

79.841 

184.544 

264.385 

167.620 

37 

83.333 

217.133 

79.841 

196.902 

276.743 

175.455 

38 

83.333 

225.595 

79.841 

203 . 952 

283 . 793 

179.925 

39 

79 . 850 

225.595 

77.434 

203.952 

281.386 

177.399 

40  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  15. — Measurements  in  Centimeters  of  Steers  at  Beginning 
of  Summer  Periods  and  End  of  Winter  Periods 


STEER  585 


5-22-14 

4-17-15 

4-12-16 

4-7-17 

5-2-18 

4-27-19 

4-21-20 

Height  at  withers 

79.5 

91.0 

103.5 

114.5 

121.5 

124.0 

128.0 

Height  at  hips 

83.5 

93.0 

107.0 

117.0 

126.5 

128.0 

130.0 

Girth  of  throat 

47.0 

52.0 

64.0 

72.0 

79.0 

78.0 

85.0 

Depth  of  chest 

32.5 

41.0 

49.5 

55.0 

60.0 

61.0 

62.5 

Width  of  chest 

18.0 

21.0 

23.0 

25.0 

26.0 

29.0 

30.0 

Width  of  paunch 

20.4 

31.0 

38.0 

42.0 

45.5 

52.5 

54.5 

Foreleg  elbow  to  ground 

49.0 

53.0 

63.0 

65.0 

72.0 

72.0 

74.0 

Point  of  shoulder  to  top  hip  point . . . 

60.0 

68.0 

87.0 

94.0 

96.0 

101.5 

109.0 

Point  of  shoulder  to  ground 

60.0 

63.0 

73.0 

77.0 

81.5 

85.0 

92.0 

Poll  to  point  of  muzzle 

26.0 

31.0 

39.0 

44.0 

49.0 

49.0 

49.0 

Heart  girth 

85.0 

102.0 

121.0 

140.0 

149.0 

153.0 

163.0 

Paunch  girth 

85.0 

124.0 

139.0 

157.0 

162.0 

183.0 

192.0 

Width  of  hips 

20.0 

27.0 

34.0 

39.0 

42.5 

43.0 

46.0 

Width  of  loin 

14.0 

18.5 

21.5 

24.5 

25.0 

32.5 

31.5 

STEER  528 


Date 

6-11-14 

4-17-15 

4-12-16 

4-7-17 

5-2-18 

4-27-19 

4-21-20 

Height  at  withers 

81.8 

109.0 

122.5 

130.0 

138.5 

140.0 

143.0 

Height  at  hips 

86.3 

110.5 

126.5 

136.0 

141.0 

143.0 

144.5 

Girth  of  throat 

54.0 

75.0 

83.0 

94.0 

100.0 

96.0 

100.0 

Depth  of  chest 

36.5 

53.0 

61.5 

66.0 

73.0 

76.0 

78.0 

Width  of  chest 

20.3 

30.0 

33.5 

40.0 

41.0 

41.0 

43.5 

Width  of  paunch 

25.0 

41.0 

46.0 

59.0 

61.0 

61.0 

64.0 

Foreleg  elbow  to  ground 

50.5 

62.0 

70.0 

75.0 

77.0 

79.5 

80.0 

Point  of  shoulder  to  top  hip  point . . . 

64.0 

87.0 

103.0 

112.0 

114.0 

122.0 

125.0 

Point  of  shoulder  to  ground 

60.5 

72.0 

82.0 

86.0 

91.0 

92.0 

96.0 

Poll  to  point  of  muzzle 

27.5 

38.0 

45.0 

54.0 

56.0 

57.0 

58.0 

Heart  girth 

95.0 

136.0 

157.0 

175.0 

191.0 

194.5 

201.0 

Paunch  girth 

98.0 

144.0 

166.0 

193.0 

206.0 

209.0 

217.0 

Width  of  hips 

22.7 

35.0 

42.5 

49.5 

54.5 

56.0 

59.0 

Width  of  loin 

15.3 

24.0 

27.0 

31.5 

35.0 

38.5 

36.0 

STEER  579 


Date 

5-30-14 

4-17-15 

4-12-16 

4-7-17 

5-2-18 

4-27-19 

4-21-20 

Height  at  withers 

81.5 

104.0 

114.5 

126.0 

135.5 

137.5 

139.0 

Height  at  hips 

75.4 

105.0 

117.0 

128.5 

136.5 

137.5 

140.0 

Girth  of  throat 

49.0 

64.0 

69.0 

73.0 

83.0 

79.0 

85.0 

Depth  of  chest 

34.5 

47.0 

54.5 

59.5 

66.0 

67.5 

69.0 

Width  of  chest 

18.5 

23.0 

25.5 

29.0 

32.0 

31.5 

36.0 

Width  of  paunch 

23.0 

32.0 

40.0 

46.0 

53.0 

49.5 

52.0 

Foreleg  elbow  to  ground 

51.0 

63.0 

68.0 

75.0 

79.0 

83.0 

84.0 

Point  of  shoulder  to  top  hip  point . . . 

62.0 

84.0 

99.0 

105.0 

114.0 

121.0 

121.0 

Point  of  shoulder  to  ground 

57.0 

68.0 

78.0 

85.0 

88.0 

92.0 

95.0 

Poll  to  point  of  muzzle 

27.5 

36.0 

43.0 

49.0 

53.0 

52.5 

53.0 

Heart  girth 

90.0 

120.0 

137.0 

150.0 

168.0 

169.0 

171 .0 

Paunch  girth 

92.0 

129.0 

142.0 

162.0 

181.0 

181.0 

187.0 

Width  of  hips 

21.5 

30.0 

36.0 

41  .0 

47.0 

47.0 

49.0 

Width  of  loin  

14.9 

21.0 

20.0 

23.5 

27.0 

31.5 

31.5 

The  Influence  of  the  Plane  of  Nutrition 


41 


Table  15  (Continued). — Measurements  in  Centimeters  of  Steers  at 
Beginning  of  Summer  Periods  and  End  of  Winter  Periods 


STEER  578 

STEER  577 

STEER 

575 

5-2-18 

4-27-19 

4-21-20 

5-2-18 

4-27-19 

4-21-20 

5-2-18 

Height  at  withers 

107.0 

113.0 

118.5 

113.5 

127.0 

134.0 

100.0 

Height  at  hips 

110.0 

114.0 

120.5 

116.75 

130.0 

136.5 

101.5 

Girth  of  throat 

64.0 

63.0 

75.0 

73.0 

79.0 

90.0 

60.0 

Depth  of  chest 

51.0 

53.5 

57.5 

56.0 

64.5 

71.5 

46.5 

"Width  of  chest 

24.0 

25.0 

27.5 

28.0 

32.5 

37.0 

21.0 

Width  of  paunch 

39.5 

46.0 

55.0 

40.0 

52.0 

55.0 

36.5 

Foreleg  elbow  to  ground 

64.5 

67.0 

69.0 

70.0 

76.5 

79.0 

60.0 

Point  of  shoulder  to  top  hip  point . . . 

86.0 

91.5 

97.0 

89.0 

106.0 

109.0 

78.0 

Point  of  shoulder  to  ground 

72.0 

76.5 

82.0 

78.0 

88.0 

95.0 

69.0 

Poll  to  point  of  muzzle 

40.0 

42.0 

44.0 

43.5 

50.0 

53.0 

38.5 

Heart  girth 

128.0 

132.0 

145.0 

141.0 

165.0 

180.0 

119.0 

Paunch  girth 

146.0 

162.0 

183.0 

149.0 

179.0 

197.0 

131.0 

Width  of  hips 

34.0 

35.5 

41.0 

34.5 

41.5 

46.5 

27.5 

Width  of  loin 

19.0 

24.0 

26.0 

23.5 

31.0 

35.0 

18.0 

STEER  575  (Cont.) 

STEER  574 

STEER  573 

Date 

4-27-19 

4-21-20 

5-2-18 

4-27-19 

4-21-20 

5-2-18 

4-27-19 

Height  at  withers 

106.5 

114.0 

99.0 

104.0 

111.0 

102.0 

109.0 

Height  at  hips 

109.0 

116.5 

103.0 

107.0 

114.5 

102.5 

109.5 

■Girth  of  throat 

63.0 

03.0 

63.0 

65.5 

72.0 

66.0 

68.0 

Depth  of  chest 

52.0 

55.0 

49.0 

52.5 

57.5 

52.5 

56.5 

Width  of  chest 

25.0 

27.5 

23.5 

26.0 

25.5 

26.0 

27.0 

Width  of  paunch 

43.0 

44.5 

37.0 

42.0 

42.5 

39.0 

44.5 

Foreleg  elbow  to  ground 

64.0 

70.0 

59.0 

64.0 

68.0 

58.0 

63.0 

Point  of  shoulder  to  top  hip  point . . . 

87.0 

94.0 

80.0 

87.0 

96.0 

77.0 

87.0 

Point  of  shoulder  to  ground 

72.0 

79.0 

66.5 

71.0 

76.0 

68.0 

71.0 

Poll  to  point  of  muzzle 

42.0 

47.0 

39.0 

42.5 

46.0 

40.0 

43.0 

Heart  girth 

130.0 

142.0 

127.0 

132.0 

145.0 

131.0 

137.0 

Paunch  girth 

148.0 

159.0 

135.0 

144.0 

154.0 

141.0 

153.0 

Width  of  hips 

31.0 

36.0 

30.0 

33.0 

36.0 

31.0 

33.5 

Width  of  loin 

22.0 

23.0 

21.0 

22.0 

23.0 

20.0 

25.0 

STEER  573  (Cont.) 

STEER  572 

STEER  571 

Date 

4-21-20 

5-2-18 

4-27-19 

4 21-20 

5-2-18 

4-27-19 

1-21-20 

Height  at  withers 

117.0 

98.5 

105.0 

107.5 

100.25 

114.5 

118.5 

Height  at  hips 

115.5 

99.25 

105.4 

110.5 

104.75 

117.5 

123.0 

Girth  of  throat 

74.0 

60.0 

62.0 

67.0 

65.0 

75.5 

80.0 

Depth  of  chest 

61.0 

46.0 

48.5 

51.0 

51.0 

57.5 

62.5 

Width  of  chest 

28.5 

23.0 

24.0 

26.0 

26.5 

31.5 

33.0 

Width  of  paunch 

47.0 

33.5 

40.0 

47.0 

40.0 

47.5 

50.0 

Foreleg  elbow  to  ground 

70.0 

61.0 

62.0 

69.0 

60.5 

69.0 

73.0 

Point  of  shoulder  to  top  hip  point.  . . 

90.0 

76.0 

83.5 

89.0 

81.0 

98.0 

101.0 

Point  of  shoulder  to  ground 

77.0 

69.0 

71.5 

76.5 

67.5 

77.5 

81.0 

Poll  to  point  of  muzzle 

46.0 

39.0 

41.5 

44.0 

39.0 

47.0 

49.0 

Heart  girth 

155.0 

116.0 

124.0 

133.0 

131.0 

149.0 

160.0 

Paunch  girth 

166.0 

124.0 

136.0 

158.0 

143.0 

162.5 

175.0 

Width  of  hips 

37.0 

29.25 

31.0 

34.0 

31.5 

38.0 

41.0 

Width  of  loin 

27.5 

18.25 

19.5 

27.0 

19.75 

23.0 

28.0 

42  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  16. — Measurements  in  Centimeters  of  Steers  at  End  of 
Summer  Periods  and  Beginning  of  Winter  Periods 


STEER  585 


Date 

10-19-14 

10-13-15 

10-8-16 

10-3-17 

10-28-18 

10-23-19 

10-17-20 

Height  at  withers 

87.5 

94.7 

109.0 

118.5 

122.0 

125.0 

129.5 

Height  at  hips 

92.0 

100.0 

114.0 

122.0 

126.0 

127.5 

130.5 

Girth  of  throat 

52.0 

53.0 

64.0 

70.0 

73.0 

79.0 

81.0 

Depth  of  chest 

39.5 

42.0 

50.0 

55.0 

59.0 

61.0 

65.5 

Width  of  chest 

19.0 

19.0 

23.0 

25.0 

25.0 

29.5 

36.0 

Width  of  paunch 

29.5 

33.5 

40.0 

43.5 

44.0 

52.0 

61.0 

Foreleg,  elbow  to  ground.  . . . 

52.0 

59.0 

65.0 

69.0 

73.0 

74.0 

74.0 

Point  of  shoulder  to  top  hip 
point 

68.5 

73.0 

91.0 

91.0 

99.0 

102.0 

114.0 

Point  of  shoulder  to  ground. . 

64.0 

64.0 

75.0 

81 .0 

82.0 

85.5 

89.0 

Poll  to  point  of  muzzle 

32.0 

33.0 

43.0 

46.5 

48.0 

48.5 

51.0 

Heart  girth 

99.0 

107.0 

123.0 

139.0 

145.0 

154.0 

169.0 

Paunch  girth 

116.0 

129.0 

145.0 

160.0 

166.0 

185.0 

204.0 

Width  of  hips 

26.5 

29.7 

36.0 

38.0 

41.0 

43.5 

47.5 

Width  of  loin 

17.0 

18.0 

20.0 

24.0 

26.0 

30.0 

36.0 

STEER  528 


Date 

10-19-14 

10-13-15 

10-8-16 

10-3-17 

10-28-18 

10-29-19 

10-17-20 

Height  at  withers 

101.5 

115.5 

127.0 

133.5 

139.0 

144.0 

144.0 

Height  at  hips 

103.5 

120.5 

131.5 

138.5 

143.0 

144.5 

143.0 

Girth  of  throat 

68.0 

73.0 

83.0 

93. 

95.0 

99.0 

102.0 

Depth  of  chest 

46.5 

55.5 

64.5 

71.0 

73.5 

78.0 

79.0 

Width  of  chest 

26.0 

28.0 

35.5 

38.5 

38.0 

41.0 

46.0 

Width  of  paunch 

39.5 

46.7 

52.5 

55.5 

55.0 

62.5 

65.0 

Foreleg,  elbow  to  ground .... 

60.0 

70.0 

72.0 

76.0 

81.0 

81.5 

77.0 

Point  of  shoulder  to  top  hip 
point 

80.0 

95.0 

104.0 

113.0 

122.0 

124.5 

120.0  . 

Point  of  shoulder  to  ground. . 

67.0 

77.0 

87.0 

88.0 

89.0 

94.0 

96.0 

Poll  to  point  of  muzzle 

35.0 

40.0 

50.0 

55.5 

55.0 

57.0 

58.0 

Heart  girth 

125.0 

144.0 

165.0 

180.0 

189.0 

198.5 

205.0 

Paunch  girth 

144.0 

164.0 

183.0 

192.0 

198.0 

214.0 

222.0 

Width  of  hips 

32.0 

39.2 

46.5 

51.0 

54.5 

59.0 

60.0 

Width  of  loin 

20.5 

23.0 

30.0 

31.0 

34.5 

36.5 

39.5 

STEER  579 


Date  

10-19-14 

10-13-15 

10-8-16 

10-3-17 

10-28-18 

10-29-19 

10-17-20 

Height  at  withers 

96.0 

110.0 

121.0 

129.5 

135.5 

138.5 

139.5 

Height  at  hips 

101.5 

114.0 

125.0 

133.5 

137.0 

138.5 

141.0 

Girth  of  throat 

59.0 

62.0 

69.0 

80.0 

77.0 

78.0 

81.5 

Depth  of  chest 

42.0 

49.5 

55.0 

61.5 

64.0 

65.5 

68.5 

Width  of  chest 

21.0 

24.0 

29.0 

30.5 

29.5 

31.5 

39.0 

Width  of  paunch 

31.5 

39.0 

43.5 

45.5 

48.0 

47.5 

52.0 

Foreleg,  elbow  to  ground .... 
Point  of  shoulder  to  top  hip 

60.0 

67.5 

69.0 

74.0 

82.5 

83.0 

78.0 

point 

79.0 

91.0 

100.0 

112.0 

120.0 

119.0 

118.0 

Point  of  shoulder  to  ground. . 

65.0 

75.0 

85.0 

85.0 

90.0 

90.5 

94.5 

Poll  to  point  of  muzzle 

34.0 

38.0 

46.0 

51.5 

53.0 

52.0 

54.0 

Heart  girth 

110.5 

124.0 

143.0 

155.0 

163.0 

164.0 

176.5 

Paunch  girth 

124.0 

143.0 

155.0 

167.0 

172.0 

176.0 

191.0 

Width  of  hips 

28.0 

32.7 

39.0 

43.5 

47.0 

48.0 

50.0 

Width  of  loin 

20.0 

20.5 

22.0 

25.0 

26.5 

30.5 

31.5 

The  Influence  of  the  Plane  of  Nutrition 


43 


Table  16  (Continued). — Measurements  in  Centimeters  of  Steers  at  End  of 
Summer  Periods  and  Beginning  of  Winter  Periods 


STEER  678 

STEER  577 

Date 

11-3-17 

10-28-18 

10-29-19 

10-17-20 

11-3-17 

10-28-18 

10-29-19 

Height  at  withers 

97.0 

110.5 

113.5 

121.0 

97.0 

122.0 

132.0 

Height  at  hips 

100.0 

111.5 

117.0 

123.0 

102.0 

124.5 

135.0 

Girth  of  throat 

56.0 

60.0 

66.0 

69.0 

60.0 

73.0 

79.0 

Depth  of  chest 

43.0 

50.5 

53.5 

59.0 

45.0 

59.0 

67.0 

Width  of  chest* 

21.5 

24.5 

24.0 

27.5 

27.0 

30.0 

35.5 

Width  of  paunch 

40.5 

43.0 

49.5 

51.0 

39.0 

46.0 

52.5 

Foreleg,  elbow  to  ground  .... 

57.0 

64.0 

69.0 

69.0 

61.0 

71.0 

80.0 

Point  of  shoulder  to  top  hip, 

point 

76.0 

90.0 

94.0 

105.0 

74.0 

100.0 

109.0 

Point  of  shoulder  to  ground . . 

67.0 

75.0 

81.0 

82.0 

68.0 

81.0 

89.5 

Poll  to  point  of  muzzle 

35.5 

41.0 

43.0 

48.0 

35.5 

46.0 

50.5 

Heart  girth 

111.0 

126.0 

137.0 

151.0 

115.0 

150.0 

171.0 

Paunch  girth 

138.0 

154.0 

171.0 

177.0 

137.0 

166.0 

186.0 

Width  of  hips 

29.0 

34.0 

37.5 

41.0 

27.5 

38.0 

44.0 

Width  of  loin 

17.5 

20.5 

24.0 

26.5 

18.5 

25.0 

33.0 

STEER  577— (C  ont  .) 

STEER  575 

STEER  574 

Date 

10-17-20 

11  3-17 

10-28-18 

10-29-19 

10-17-20 

11-3-17 

10-28-18 

Height  at  withers 

136.5 

91.5 

102.5 

111.0 

118.5 

91.0 

102.0 

HeiTht  at  hips 

138.0 

95.0 

104.5 

115.5 

119.0 

95.5 

105.0 

Girth  of  throat 

89.0 

55.0 

57.0 

70.0 

72.0 

58.5 

57.0 

Depth  of  chest 

73.0 

42.0 

47.5 

54.0 

57.5 

44.5 

49.5 

Width  of  chest 

40.0 

21.0 

22.0 

26.0 

29.5 

23.0 

24.0 

Width  of  paunch 

56.0 

36.0 

42.5 

47.5 

49.5 

39.0 

40.0 

Foreleg,  elbow  to  ground  .... 

81.5 

57.0 

62.0 

69.0 

70.0 

56.0 

60.0 

Point  of  shoulder  to  top  hip 

point 

122.0 

72.0 

85.0 

91.5 

100.0 

72.0 

83.0 

Point  of  shoulder  to  ground. . 

93.0 

65.0 

69.0 

78.5 

79.0 

63.0 

67.0 

Poll  to  point  of  muzzle 

56.0 

34.0 

40.0 

45.0 

50.0 

36.0 

41.0 

Heart  girth 

181.0 

107.0 

119.0 

137.0 

146.0 

113.0 

128.0 

Paunch  girth 

201.0 

129.0 

148.0 

165.0 

169.0 

132.0 

148.0 

Width  of  hips 

48.0 

24.0 

28.0 

33.0 

37.5 

27.0 

30.5 

Width  of  loin 

35.5 

16.5 

18.5 

23.0 

26.5 

17.0 

22.0 

STEER 

STEER  574— (Cont.) 

STEER  573 

572 

Date 

10-29-19 

10-17-20 

11-3-17 

10-28-18 

10-29-19 

10-17-20 

11-3-17 

Height  at  withers 

108.5 

114.0 

92.0 

106.5 

114.5 

119.0 

89.5 

Height  at  hips 

112.0 

116.5 

94.0 

107.0 

115.0 

118.5 

93.0 

Girth  of  throat 

69.0 

70.0 

58.0 

64.0 

74.0 

75.0 

55.0 

Depth  of  chest 

56.0 

60.5 

44.5 

51.5 

58.0 

68.0 

40.5 

Width  of  chest 

29.0 

32.5 

24.5 

25.0 

30.0 

30.5 

22.5 

Width  of  paunch 

46.5 

47.0 

39.0 

43.0 

48.5 

50.5 

36.0 

Foreleg,  elbow  to  ground  .... 

66.0 

65.0 

55.0 

62.0 

68.0 

66.5 

56.0 

Point  of  shoulder  to  top  hip 

point 

92.5 

100.0 

71.0 

83.0 

94.0 

101.0 

70.0 

Point  of  shoulder  to  ground. . 

75.0 

78.0 

63.0 

72.0 

75.5 

79.0 

62.0 

Poll  to  point  of  muzzle 

45.5 

49.0 

35.0 

41.0 

44.5 

49.0 

34.0 

Heart  girth 

145.0 

150.0 

116.0 

133.0 

149.0 

159.0 

116.0 

Paunch  girth 

160.0 

165.0 

138.0 

150.0 

167.0 

175.0 

126.0 

Width  of  hips 

35.0 

38.0 

28.0 

31.5 

36.0 

39.0 

25.0 

Width  of  loin 

25.0 

27.5 

19.0 

21.0 

28.0 

27.5 

16.5 

44  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  16  (Continued). — Measurements  in  Centimeters  of  Steers  at  End  of 
Summer  Periods  and  Beginning  Winter  Periods 


STEER  572— (Cont.) 

STEER  671 

10-28-18 

102.25 

10-29-19 

107.5 

10-17-20 

11-3-17 

10  28-18 
107.0 

10-29-19 

116.0 

10-17-20 

120.0 

Height  at  withers 

115.5 

83.5 

Height  at  hips 

102.25 

109.0 

113.0 

88.5 

111.0 

120.5 

123.5 

Girth  of  throat 

58.0 

65.0 

65.0 

54.0 

68.0 

72.0 

78.0 

Depth  of  chest 

46.5 

50.0 

54.5 

39.0 

53.0 

59.5 

63.5 

Width  of  chest 

23.5 

24.0 

29.0 

20.0 

29.0 

33.5 

34.5 

Width  of  paunch 

39.5 

41.0 

49.0 

35.0 

48.0 

50.0 

52.5 

Foreleg,  elbow  to  ground.  . . . 

63.0 

67.0 

67.0 

53.0 

66.0 

71.5 

72.0 

Point  of  shoulder  to  top  hip 
point 

82.0 

85.5 

98.0 

65.0 

88.0 

101.0 

110.0 

Point  of  shoulder  to  ground . . 

67.0 

76.0 

77.0 

57.0 

70.0 

81.5 

80.0 

Poll  to  point  of  muzzle 

39.0 

42.0 

46.0 

32.0 

42.0 

49.0 

51.0 

Heart  girth 

120.0 

129.0 

137.0 

100.0 

140.0 

156.0 

165.0 

Paunch  girth. 

138.0 

145.0 

163.0 

125.0 

163.0 

167.0 

182.0 

Width  of  hips 

30.0 

32.0 

35.0 

24.0 

35.0 

39.5 

43.0 

Width  of  loin 

18.5 

23.0 

26.5 

16.0 

23.0 

29.0 

28.5 

Table  17. — Measurements*  in  Centimeters  of  Control  Animals 
at  Time  of  Slaughter 


Steer  No 

554 

552 

523 

526 

512 

531 

525 

509 

Height  at  withers  . 

90.5 

96.5 

128.8 

130.0 

153.0 

114.75 

124.8 

140.0 

Height  at  hips .... 

96.8 

98.0 

130.0 

140.5 

150.5 

115.5 

126.8 

139.0 

Girth  of  throat. . . . 

55.0 

60.0 

87.0 

94.0 

100.0 

72.0 

80.0 

90.5 

Depth  of  chest .... 

38.5 

44.0 

66.0 

71.5 

80.0 

54.5 

62.0 

67.0 

Width  of  ohest .... 

22.3 

25.0 

36.0 

43.0 

44.5 

26.5 

30.5 

40.5 

Width  of  Paunch. . 
Foreleg,  elbow  to 

25.6 

33.3 

62.0 

57.0 

62.0 

42.0 

54.0 

53.0 

ground 

Point  of  shoulder  to 

60.0 

61.2 

76.5 

85.0 

88.0 

72.5 

73.8 

84.5 

top  hip  point  . . . 
Point  of  shoulder  to 

70.5 

75.0 

112.0 

124.0 

127.0 

94.0 

105.0 

116.0 

ground 

Poll  to  point  of 

67.0 

70.0 

88.0 

94.0 

101.0 

88.5 

85.0 

98.5 

muzzle 

31.5 

35.0 

52.5 

54.0 

55.0 

43.0 

51.0 

52.0 

Heart  girth 

101.0 

114.0 

175.0 

197.0 

210.0 

146.0 

160.0 

186.0 

Paunch  girth 

103.0 

125.0 

210.0 

212.0 

220.0 

160.0 

189.0 

204.0 

Width  of  hips 

23.1 

27.0 

46.0 

53.0 

56.0 

35.5 

40.5 

48.5 

Width  of  loin 

17.0 

21.7 

37.0 

41.0 

43.5 

27.5 

31.5 

41.0 

*From  unpublished  data  furnished'  by  C.  R.  Moulton,  department  agricultural  ohemistry, 
Missouri  Agricultural  Experiment  Station. 


The  Influence  on  the  Plane  of  Nutrition 


45 


Table  18. — Composition  of  Control  Animals 


Steer 

Age  days 

Weight  pounds 

Composition  of  body 

Dry  matter 
per  cent 

Protein 
per  oent 

Fat 

per  cent 

554 

90 

196.  0 

32 . 836 

20.038 

7.395 

552 

160 

256.2 

35.928 

19.469 

10.555 

523 

798 

864.2 

39.479 

19.219 

15.134 

526 

1217 

1088.2 

44.552 

18.813 

20.435 

512 

1454 

1250.4 

48.001 

18.094 

24.299 

531 

588 

479.6 

37.227 

20.263 

10.355 

525 

800 

694.6 

37.338 

20.031 

11.787 

509 

1363 

1004.2 

42.079 

20.181 

16.232 

From  unpublished  data  furnished  by  C.  R.  Moulton,  department  agricultural  chemistry 
Missouri  Agricultural  Experiment  Station. 


Table  19. — Energy  Value  of  Gains,  Calculated  for  Summer  Ferioes 


Periodt 

Steer 

1 

Therms 

2 

Therms 

3 

Therms 

4 

Therms 

5 

Therms 

6 

Therms 

7 

Therms 

Group  1 

528 

577 

.95575 

1.0918 

1.0918 

1.2279 

1.2279 

1.7136 

1.7136 

2.1993 

2.500 

3.000 

571 

1.0918 

1.2279 

1.7136 

Group  II 

579 

578 

.95575 

1.0583 

1.0583 

1.0583 

1 . 0583 
1 . 1608 

1.1608 

1.4104 

1.5352 

1.66 

573 

1 . 0583 

1.0583 

1 . 1608 

Group  III 

585 

575 

.8343 

.9445 

.9445 
1 . 0548 

.9445 

1.1013 

1.0548 

1.1013 

1.1479 

1.649 

574 

.9445 

1.0548 

1.1013 

572 

.9445 

1.0548 

1.1013 

Table  18  shows  the  ages,  weights,  and  percentage  composition  of  live  weight  of  the  control 
animals  used  in  this  study. 

tThese  same  values  apply  also  to  the  winter  periods  of  the  seven  younger  steers.  In  the 
case  of  the  three  older  animals,  Nos.  528,  579,  and  585,  the  first  winter  period  corresponds  to  the 
second  summer  period  and  so  on,  thus  making  the  sixth  winter  period  correspond  to  the  seventh 
summer  period. 


From  these  data  the  composition  of  the  gains  was  estimated  for  all  periods  except  period 
seven  of  528.  No  control  animal  to  fit  this  period  could  be  found.  The  value  of  three  therms  per 
pound  gain  was  used  in  this  case.  This  value  was  assumed  on  the  basis  of  Armsby’s  calculation 
(3f)  of  the  energy  value  of  gains  from  Lawes  and  Gilbert’s  analyses  on  four-year-old  fattening 
cattle. 

Table  19  shows  the  estimated  energy  value  of  the  gains  by  periods  for  all  the  steers. 

Table  20  shows  distribution  of  the  control  animals.  Check  animals  were  first  fitted  to  the 
old  steer  in  eaoh  group;  then  the  young  animals  were  oompared  with  the  old  animal  in  their  re- 
spective groups  rather  than  with  check  animals  direct. 


46  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Table  20. — Distribution  oe  Control  Animals. 


Period 

1 

2 

3 

4 

5 

6 

7 

Steer 

528 

552 

Inter- 

polate 

523 

Inter- 

polate 

526 

512 

577 

571 

Period 
2 of 
528 

523 

Period 
4 of 
528 

579 

552 

I nter- 
polate 

Inter- 

polate 

525 

Inter- 

polate 

Inter- 

polate 

509 

578 

573 

Period 
2 of 
579 

Period 
3 of 
579 

525 

585 

554 

Inter- 

polate 

Inter- 

polate 

531 

Inter- 

polate 

525 

523 

575 

574 

572 

Period 
3 of. 
585 

531 

Period 
5 of 
585 

Fig.  1. — Weights — Comparison  of  control  animals  and  experimental  animals. 


The  Influence  on  the  Plane  of  Nutrition 


47 


Fig.  2. — Heart  Girth — Comparison  of  control  animals  and  experimental  animals. 


Fig-  3. — Depth  of  Chest — Comparison  of  control  animals  and  experimental  animals. 


48  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


Fig.  5. — Height  • at  Withers — Comparison  of  control  animals  and  experimental  animals. 


UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 

AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  52 


SCARRED  ENDOSPERM  AND 
SIZE  INHERITANCE 


ERRATA 

transpose  table  headings  of  the  two 
(ties  opposite  page  6. 

Page  7,  table  3:  Strike  out  “Average  Ker- 
Weight  in  mgs.”  in  box  head  of  last  col- 

!n. 


5 OF  MAIZE 


rized  June  1,  1922.) 


COLUMBIA,  MISSOURI 

JULY,  1922 


48  Missouri  Agr.  Exp.  Station  Research  Bulletin  51 


O — f\>  to.  -C*  cn  O'  CD  <£>  o — rj  to  K 0>  0'  3 CO 


Fig.  5. — Height  at  Withers — Comparison  of  control  animals  and  experimental  animals. 


UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 

AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  52 


SCARRED  ENDOSPERM  AND 
SIZE  INHERITANCE 
IN  KERNELS  OF  MAIZE 

(Publication  authorized  June  1,  1922.) 


COLUMBIA,  MISSOURI 

JULY,  1922 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL, 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY  P.  E.  BURTON  H.  J.  BLANTON 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 

J.  C.  JONES,  PH.  D.,  LL.  D.,  PRESIDENT  OF  THE  UNIVERSITY 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 

STATION  STAFF 

JULY,  1922 

AGRICULTURAL  CHEMISTRY 


C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  Ph.  D. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  SiEveking,  B.  S.  in  Agr. 

A.  M.  Cowan,  A.  M. 

AGRICULTURAL  ENGINEERING 

J.  C.  WoolEy,  B.  S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumeord,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

VV.  J.  Robbins,  Ph.  D. 

E.  F.  Hopkins,  Ph.  D. 

DAIRY  HUSBANDRY 
A.  C.  Ragsdale,  B.  S.  in  Agr. 

W.  W.  Swett,  A.  M. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride,  B.  S.  in  Agr. 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  A.  M. 

O.  W.  Letson,  B.  S.  in  Agr. 

Alva  C.  Hill,  B.  S.  in  Agr. 

Miss  Pearl  Drummond,  A.  A.* 


RURAL  LIFE 
O.  R.  Johnson,  A.  M. 

S.  D.  Gromer,  A.  M. 

E.  L.  Morgan,  A.M. 

Ben  H.  Frame,  B.  S.  in  Agr. 
Owen  Howells 


HORTICULTURE 
V.  R.  Gardner,  M.  S.  A. 

H.  D.  Hooker,  Jr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  M.  S. 

F.  C.  Bradford,  M.  S. 

H.  G.  Swartwout,  B.  S.  in  Agr. 

poultry  husbandry 

H.  L.  Kempster,  B.  S. 

Earl  W.  Henderson,  B.S. 

SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M 
I W.  A.  Albrecht,  Ph.  D. 

F.  L.  Duley,  A.M. 

R.  R.  Hudelson,  A.M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr 
Richard  Bradfield,  A.  B. 

VETERINARY  SCIENCE 
J.  W.  Connaway,  D.  V.  S.,  M.  D. 

L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 

OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Secretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Jane  Frodsham,  Librarian. 

E.  E.  Brown,  Business  Manager. 


In  service  of  U.  S.  Department  of  Agriculture. 


Scarred  Endosperm  and  Size  Inheri- 
tance in  Kernels  of  Maize 


William  H.  Eyster* 

In  the  summer  of  1920  the  writer  found  in  a field  of  corn  in  cen- 
tral Pennsylvania  a number  of  plants  with  striking  chlorophyl  patterns 
which  are  unlike  any  that  have  yet  been  described.  These  plants  were 
numbered  and  marked  in  the  field  so  that  they  could  be  identified  at 
harvest  time.  The  matured  ears  were  sent  to  the  writer  at  the  Univer- 
sity of  Missouri  by  Mr.  Webster  Snyder  in  whose  field  they  were  dis- 
covered. Plantings  were  made  from  each  ear  in  the  greenhouse  the 
following  winter  and  the  seedlings  were  found  to  be  entirely  green. 
One  plant  from  each  ear  was  grown  to  maturity  in  the  greenhouse  and 
self  pollinated.  In  the  summer  of  1921  field  plantings  were  made  from 
the  original  ears  and  also  from  the  self  pollinated  greenhouse  ears.  The 
F2  progenies  segregated  plants  with  the  chlorophyl  patterns  of  the 
original  plants  together  with  a number  of  other  characters,  including 
a pistillate  plant  similar  in  appearance  to  tassel  ear  (Emerson,  1920)  and 
the  endosperm  character  described  in  this  paper,  which  has  been  desig- 
nated scarred  endosperm. 

The  field  plantings  of  1921  were  made  at  the  Missouri  Agricul- 
tural Experiment  Station  as  part  of  a project  in  the  genetics  of  maize 
carried  on  in  the  Department  of  Field  Crops. 

DESCRIPTION  OF  SCARRED  ENDOSPERM 

Maize  kernels  with  scarred  endosperm  can  usually  be  recognized 
on  the  ear,  even  though  the  kernels  are  so  closely  arranged  that  only 
the  crowns  are  visible.  The  scarred  kernels  are  not  so  large  as  the 
normal  kernels  on  the  same  ear  and  are  commonly  pinched  ofif  so  that 
they  are  somewhat  similar  in  appearance  to  kernels  with  “rough  Men- 
tation”. 

The  scarred  character  can  more  easily  and  certainly  be  recognized 
upon  examination  of  the  abgerminal  surface  of  the  kernel.  Its  external 
appearance  is  that  of  a scar  left  after  the  healing  up  of  a deep  wound. 


•Assistant  Professor  of  Botany,  Department  of  Botany,  University  of  Missouri,  as- 
sociated with  the  Department  of  Field  Crops,  Missouri  Agricultural  Experiment  Station,  in 
the  genetic  studies  of  corn,  carried  on  by  this  Department. 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  52 


In  Fig.  1 are  shown  camera  lucida  drawings  of  a number  of  kernels 
with  scarred  endosperm.  In  Fig.  2 are  similar  drawings  of  two  scarred 
kernels  from  which  the  pericarp  overlying  the  abgerminal  surface  was 
removed.  The  nature  of  this  endosperm  character  can  best  be  seen 
from  the  drawings.  It  is  an  irregular  cavity  in  the  endosperm  on  the 
abgerminal  side  of  the  kernel.  The  cavity  consists  in  a crater-like  ex- 
cavation near  the  crown  with  divergent  and  often  branched  furrows 
extending  towards  the  base  of  the  kernel. 

The  pericarp  over  the  crater-like  excavation  near  the  crown  nearly 
always  collapses  and  causes  the  kernels  to  have  a rough  indentation. 
Occasionally  a kernel  is  found  with  the  pericarp  over  the  crater  of  the 
cavity  in  the  form  of  a blister. 

Scarred  Endosperm  and  Size  of  Kernel. — Scarred  kernels  are 
uniformly  smaller  than  normal  kernels.  In  Fig.  3 is  shown  a crown 
view  of  a series  of  representative  normal  kernels  (upper  row)  and 
scarred  kernels  (lower  row)  which  were  taken  from  the  same  ear. 
In  Fig  4 is  shown  the  same  series  of  kernels,  as  in  Fig.  3,  but  from  the 
side.  These  figures  show  in  a general  way  the  relative  differences 
in  size  between  normal  and  scarred  kernels  of  maize. 

Scarred  Endosperm  and  Thickness  of  Kernel. — The  most  con- 
spicuous size  difference  between  normal  and  scarred  kernels  is  in  the 
thickness  of  the  kernels.  Thickness  here  refers  to  the  distance  between 
the  germinal  and  abgerminal  surfaces  of  the  kernel.  The  thickness 
of  each  kernel  of  individual  ears  segregating  normal  and  scarred  ker- 
nels was  measured  by  using  a sliding  caliper  rule  and  tabulated  as  shown 
in  Table  1.  Readings  were  made  to  the  nearest  one-half  millimeter. 
The  measurements  were  made  by  clamping  the  caliper  over  the  end 
of  the  kernel  at  a uniform  distance  from  the  crown. 

The  distributions  given  in  Table  1 show  that  for  each  ear  the  nor- 
mal kernels  are  thicker  than  those  with  scarred  endosperm.  The  mean 
thickness  of  normal  kernels  from  individual  ears  varies  from  3.856  to 
5 .722  millimeters.  The  mean  thickness  of  the  scarred  kernels  from 
the  same  ears  varies  from  3.100  to  5.360  millimeters.  The  mean  dif- 
ference in  thickness  of  the  normal  and  scarred  kernels  from  the  ears 
studied  varied  from  0.295  to  0.823  millimeter.  The  mean  thickness 
of  the  normal  kernels  of  the  eight  distributions  listed  in  Table  1 con- 
sidered collectively  is  4.500  — 0.144  millimeters.  The  mean  thickness  of 
the  scarred  kernels  of  these  distributions  is  3.926^0.258  millimeters. 
The  normal  kernels  are  0.574  ± 0.295  millimeter  thicker  than  the  scarred 


Scarred  Endosperm  and  Size  Inheritance  in  Maize 


5 


kernels.  In  Fig.  5 are  given  curves  which  show  graphically  the  vari- 
ation in  thickness  of  the  normal  and  scarred  kernels.  These  curves 
represent  the  total  frequencies  of  the  distributions  listed  in  Table  1. 

Scarred  Endosperm  and  Weight  of  Kernel. — The  kernels  of 
each  ear  studied  were  weighed  individually  to  the  nearest  milligram 
and  tabulated  as  shown  in  Table  2.  In  every  case  the  mean  weight 
of  the  normal  kernels  is  higher  than  the  mean  weight  of  the  scarred 
kernels  from  the  same  ear.  The  mean  weights  of  the  normal  kernels 
from  the  ears  studied  varied  from  251.92  to  341.10  milligrams.  The 
mean  weights  of  the  scarred  kernels  from  the  same  ears  varied  from 
232.70  to  329.60  milligrams.  The  differences  in  the  means  of  the  in- 
dividual ears  varied  from  1.24  milligrams  for  ear  1243-2  to  19.13  milli- 
grams for  ear  1238-16.  In  order  to  obtain  a general  expression  of  the 
mean  weights  of  the  normal  and  scarred  kernels  the  distributions  in 
Table  2 are  considered  collectively.  The  mean  weight  of  the  normal 
kernels  from  the  eight  ears  is  274  milligrams  and  the  mean  weight  of 


Fig.  5. — Variation  in  thickness  of  normal  and  scarred  kernels. 


6 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  52 


the  scarred  kernels  is  259.57  milligrams.  This  is  a difference  of  14.43 
— 1.29  milligrams.  In  Fig.  6 are  curves  of  variation  in  weight  of  nor- 
mal and  scarred  kernels  when  the  eight  distributions  of  Table  2 are 
considered  collectively.  In  many  respects  these  curves  are  similar  to 
those  for  thickness  of  kernel  given  in  Fig.  5. 

The  normal  and  scarred  kernels  respectively  of  each  ear  were 
weighed  en  masse  with  the  results  given  in  Table  3.  From  these  total 
weights  average  kernel  weights  were  obtained  that  do  not  involve  the 
errors  due  to  the  separate  weighing  of  the  individual  kernels.  The  av- 
erage kernel  weights  are  in  fairly  close  agreement  with  the  mean  weights 
as  given  in  Table  2. 

The  normal  and  scarred  kernels  from  ear  1243-2  differ  only  slightly 
in  average  thickness  and  have  approximately  the  same  kernel  weight. 
The  mean  weight  of  the  normal  kernels  is  given  in  Table  2 as  1.24 
milligrams  greater  than  that  of  the  scarred  kernels.  The  average 
weight,  however,  of  the  normals  was  found  to  be  1.36  milligrams  less 
than  the  average  weight  of  the  scarred  kernels.  For  the  other  ears  the 
average  weight  of  the  normal  kernels  varies  from  2.17  to  20.63  milli- 
grams heavier  than  the  scarred  kernels  taken  from  the  same  ear.  The 


Fig.  6. — Variation  in  weight  of  normal  and  scarred  kernels. 


Numbei 

Descripi 

90 

1 

450 

Total 

1 Mean 

Difference 

1238-16 

Normal 

1 



384 

288.07 

Scarred 

— 

— 

139 

268.94 

19.13 

1243-2 

Normal 



424 

233.94 

Scarred 

— 

— 

145 

232.70 

1.24 

1243-8 

Normal 

1 

340 

312.12 

Scarred 

— 

— 

121 

300.17 

11.95 

1243-13 

Normal 



301 

260.86 

Scarred 

— 

— 

128 

249.84 

11.02 

1243-7 

Normal 

__ 



181 

341.10 

Scarred 

— 

— 

24 

329.60 

11.50 

1245-6 

Normal 



422 

251.92 

Scarred 

— 

— 

128 

23703 

14.89 

1245-7 

Normal 



58 

287.93 

Scarred 

— 

— 

25 

284.82 

3.11 

1245-8 

Normal 



85 

255.65 

Scarred 

— 

— 

36 

247.22 

8.43 

Total 

Normal 

1 

1 

2195 

274.00 

Scarred 

-- 

— 

746 

259.57 

14.43 

Table  2. — F2  Kernels  of  the  Cross  Normal  X Scarred 
Thickness  of  Kernels  in  Millimeters 


Pedigree 


Number 

Description 

1.0 

1.5 

2.0 

2.5 

3.0 

3.5 

4.0 

4.5 

5.0 

5.5 

6.0 

6.5 

7.0 

7.5 

Total 

Mean 

Difference 

1238-16 

Normal  

__ 

1 

4 

10 

43 

93 

77 

73 

32 

27 

17 

10 

387 

4.637 

.572 

Scarred  

- 

1 

5 

5 

11 

28 

32 

25 

17 

8 

5 

1 

- 

- 

138 

4.065 

1243-2 

Normal  

1 

28 

95 

167 

54 

37 

23 

11 

3 

419 

4.123 

.332 

Scarred  

- 

1 

2 

5 

28 

41 

39 

12 

7 

6 

5 

146 

3.791 

1243-6 

Normal  

3 

12 

26 

30 

15 

17 

29 

23 

2 

157 

5.070 

.815 

Scarred  

- 

- 

- 

1 

3 

9 

13 

7 

10 

4 

1 

1 

- 

49 

4.255 

1243-8 

Normal  

5 

23 

69 

62 

69 

62 

41 

12 

3 

346 

4.860 

.484 

Scarred  

- 

- 

4 

5 

5 

10 

26 

26 

19 

13 

9 

117 

4.376 

1243-13 

Normal  

1 

13 

43 

86 

72 

49 

16 

14 

9 

3 

__ 

306 

4.433 

.823 

Scarred  

- 

- 

11 

13 

25 

25 

15 

14 

13 

6 

1 

123 

3.610 

1245-6 

Normal  

3 

104 

118 

102 

33 

22 

18 

9 

6 

3 

418 

3.856 

.756 

Scarred 

"I 

6 

15 

29 

32 

27 

5 

4 

3 

2 

1 

125 

3.100 

1245-7 

Normal  

3 

4 

2 

15 

15 

19 

8 

2 

3 

71 

5.655 

.295 

Scarred  _ 

1 

1 

4 

6 

7 

0 

4 

2 

25 

5.360 

1245-8 

Normal  

1 

1 

7 

14 

19 

23 

18 

5 

__ 

88 

5.722 

.379 

Scarred  

1 

3 

12 

11 

6 

2 

- 

- 

35 

5.343 

Total 

Normal 

1 

9 

163 

338 

548 

337 

294 

202 

173 

96 

28 

3 

2192 

4.500 

Scarred  _ — 

"I 

~8 

37 

58 

104 

141 

132 

95 

87 

57 

28 

8 

2 

— 

758 

3.926 

.574 

Scarred  Endosperm  and  Size  Inheritance  in  Maize 


7 


data  in  Table  3 show  a greater  difference  in  weight  of  normal  and 
scarred  kernels  for  some  ears  than  the  data  in  Table  2. 


Table  3. — F2  Kernels  From  the  Cross  Normal  X Scarred 


Noi 

-mal  Kerr 

els 

Sea 

rred  Kerr 

lels 

Difference 

Total 

Average 

Total 

Average 

Average 

Pedigree 

Num- 

weight 

kernel 

Num- 

weight 

kernel 

kernel 

Number 

ber 

in  mgs. 

weight 
in  mgs. 

ber 

in  mgs. 

weight 
in  mgs. 

weight 
in  mgs. 

1238-16 

387 

108880 

281.34 

138 

37700 

273.19 

8.15 

1243-2 

442 

102325 

231.50 

124 

28875 

232.86 

—1.36 

1243-7 

185 

52200 

337.57 

25 

8200 

328.00 

9.57 

1243-8 

345 

107775 

312.10 

119 

36200 

304.20 

7.90 

1243-13 

299 

80850 

270.40 

126 

32240 

255.87 

14.53 

1245-6 

420 

89750 

213.69 

126 

24325 

193.06 

20.63 

1245-7 

60 

17650 

294.17 

25 

7300 

292.00 

2.17 

1245-8 

87 

21750 

250.00 

35 

8650 

247.14 

2.86 

Total 

2225 

581180 

261.20 

718 

183490 

255.56 

5.64 

INHERITANCE  OF  SCARRED  ENDOSPERM 

The  factor  pair  for  scarred  endosperm  is  designated  by  the  sym- 
bols Sc  sc. 

Fx  Generation. — Fx  kernels  from  the  cross  scarred  x normal , or 
its  reciprocal,  have  normal  endosperm. 

F2  Generation. — When  Fx  plants  are  self  pollinated,  ears  are 
produced  which  have  normal  and  scarred  kernels  in  ratios  approximat- 
ing 3 : 1.  In  Table  4 are  recorded  the  numbers  of  normal  and  scarred 


Table  4. — F2  Kernels  oe  the  Cross  Normal  X Scarred 


Pedigree 

Normal 

Scarred 

Total 

Ratio 

per  4 

Numbers 

kernels 

kernels 

1238-16 

390 

138 

528 

2.955 

: 1.045 

1243-2 

442 

124 

566 

3.124 

: 0.876 

1243-7 

185 

25 

210 

3.524 

: 0.476 

1243-8 

345 

119 

464 

2.974 

: 1.026 

1243-13 

299 

126 

425 

2.814 

: 1.186 

1245-6 

424 

128 

552 

3.072 

: 0.928 

1245-7 

60 

25 

85 

2.828 

: 1.172 

1245-8 

87 

35 

122 

2.853 

: 1.147 

Total  observed  2232 

720 

2952 

3.026 

: 0.974 

Total  expected  2214 

738 

2952 

3.000 

: 1.000 

Deviation  18  ± 15.88 


8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  52 


kernels  taken  from  eight  ears  of  Fx  plants  that  had  been  self  pollinated. 
The  ratios  of  the  individual  ears  vary  from  2.814  : 1.186  to  3.524  : 
0.476.  The  average  ratio  for  all  the  kernels  from  the  eight  ears  is 
3.026  : 0.974.  The  total  numbers  observed  were  2232  normal  and  720 
scarred  kernels.  This  is  a deviation  from  the  expected  distribution 
of  18  — 15.88  kernels. 

F3  Generation. — A field  planting  under  family  number  1238 
was  made  from  a self  pollinated  ear  that  segregated  kernels  with  scar- 
red endosperm.  Twenty-one  F2  plants  were  grown  to  maturity.  Three 
of  these  were  wholly  pistillate  plants.  The  remainder  were  self  pol- 
linated and  produced  ears  with  kernels  as  indicated  below : 

Kernels  all  Normal  and  All  scarred 
Normal  Scarred  kernels  Kernels 


Observed  6 9 3 

Expected  4.5  9 4.5 

Deviation  1.5  0 — 1.5 


These  numbers  are  small  but  are  in  close  agreement  with  expecta- 
tion. 

SUMMARY  AND  DISCUSSION 

Scarred  is  a new  endosperm  character  in  maize  which  consists 
in  an  irregular  cavity  in  the  endosperm  on  the  abgerminal  side  of  the 
kernel.  Kernels  with  scarred  endosperm  usually  have  a rough  indenta- 
tion. Scarred  kernels  have  been  compared  in  thickness  and  weight  with 
normal  kernels  and  it  is  evident  both  from  the  general  appearance  of 
the  kernels  and  from  the  data  given  in  this  paper  that  scarred  kernels 
are  smaller  than  the  kernels  with  normal  endosperm.  Scarred  endo- 
sperm is  inherited  as  a simple  Mendelian  recessive  character.  Cor- 
related with  scarred  endosperm  is  a difference  in  size  of  kernel  that  is 
apparently  due  to  the  same  factor. 

Emerson  and  East  (1913)  found  in  their  study  of  quantitative 
characters  in  maize  size  differences,  such  as  height  of  plant,  length 
of  ear,  and  size  of  kernel,  to  be  due  to  multiple  factors.  Such 
quantitative  characters,  however,  are  not  always  due  to  multiple  fac- 
tors. A difference  in  size,  or  in  any  quantitative  character,  between 
certain  individuals  may  be  due  to  multiple  factors,  but  a similar  size 
or  quantitative  difference  between  certain  other  individuals  may  be 
due  to  a single  factor.  Thus  differences  in  height  of  plants  are  com- 
monly due  to  a number  of  factors,  but  a large  number  of  height  dif- 


Scarred  Endosperm  and  Size  Inheritance  in  Maize 


9 


ferences  in  maize  plants  have  already  been  found  that  are  due  to  single 
factors.  As  examples  may  be  mentioned  dwarf  and  anther  ear  (Emer- 
son, R.  A.,  and  Emerson,  S.  H.,  1921),  brachytic  (Kempton,  1920)  and 
others  from  the  cultures  of  the  writer  and  other  workers  in  corn.  By 
inter  crossing  these  different  types,  progenies  can  be  produced  which 
segregate  a number  of  factors  for  height  of  plant,  and  size  inheritance 
becomes  quantitative.  The  same  principle  may  be  applied  to  other 
quantitative  differences. 

Scarred  endosperm  represents  a difference  in  size  of  kernel  which 
is  quantitative,  but  due  apparently  to  a single  factor.  In  this  re- 
spect it  is  similar  to  size  differences  in  seeds  of  beans  observed  by 
Johannsen  (1913).  In  one  of  his  pure  lines  Johannsen  found  a mu- 
tant with  a longer  seed  than  the  parent  stock.  Seed  length  of  this  mu- 
tant bean  was  found  by  Leitch  (1921)  to  be  inherited  as  a single 
Mendelian  character.  Johannsen  (1913)  also  found  in  his  cultures  a 
broad  bean  which,  when  crossed  with  the  type,  gives  an  F2  progeny  of 
1 type  : 2 intermediate  : 1 broad. 

It  is  reasonable  to  expect  that  maize  plants  will  be  found  with 
other  quantitative  differences  than  height  of  plant  and  size  of  kernel 
that  are  inherited  as  simple  Mendelian  characters. 


10 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  52 


ACKNOWLEDGMENTS 

The  writer  is  indebted  to  George  T.  Kline  for  the  drawings  in 
Figs.  1 and  2,  and  to  James  F.  Barham  for  the  photographs  in  Figs. 
3 and  4. 


LITERATURE  CITED 

Emerson,  R.  A.,  1920.  Heritable  Characters  of  Maize.  II.  Pistillate 
Flowered  Maize  Plants.  Jour.  Heredity  11  : 65-76. 

Emerson,  R.  A.  and  East,  E.  M.,  1913.  The  inheritance  of  quantita- 
tive characters  in  maize.  Neb.  Agr.  Exp.  Sta.  Research 
Bui.  2. 

Emerson,  R.  A.  and  Emerson  S.  H.,  1921.  Genetic  interrelations  of 
two  andromonoecious  types  of  maize : dwarf  and  anther 
ear  (In  manuscript). 

Johannsen,  W.,  1913.  Elemente  der  exakten  Erblichkeitslehre.  Ver- 
lag  von  Gustav  Fischer,  Jena.  Zweite  Auflage:  652-654. 

Kempton,  J.  H.,  1920.  Heritable  characters  of  maize.  III.  Brachytic 
Culms.  Jour.  Heredity  11  : 111-115. 

Leitch,  I.,  1921.  A study  of  the  segregation  of  a quantitative  char- 
acter in  a cross  between  a pure  line  of  beans  and  a mu- 
tant from  it.  Jour.  Genetics,  11  : 183-204. 


*#*• 


Fig.  1. — Maize  kernels  with  scarred  endosperm. 


Fig.  2. — Maize  kernels  with  scarred  endosperm.  The  pericarp  has  been  removed  from 
these  kernels  to  show  the  nature  of  the  scarred  endosperm. 


4.^6^  inaii 
; fe  i 8 4 I 


Fig.  3. — Kernels  with  normal  endosperm  (upper  row)  and  scarred  endosperm  (lower 

row)  from  the  same  ear. 


Fig.  4. — Kernels  with  normal  endosperm  (upper  row)  and  scarred  endosperm  (lower  row) 

from  the  same  ear. 


UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 
AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  53 


THE 

RELATION  OF  TEMPERATURE 
TO  BLOSSOMING  IN  THE 
APPLE  AND  THE  PEACH 

(Publication  Authorized  August  18,  1922) 


COLUMBIA,  MISSOURI 
AUGUST,  1922 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY  P.  E.  BURTON  H.  J.  BLANTON 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 

J.  C.  JONES,  PH.  D.,  LL.  D.,  PRESIDENT  OF  THE  UNIVERSITY 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 


STATION  STAFF 

AUGUST,  1922 

AGRICULTURAL  CHEMISTRY 


RURAL  LIFE 


O.  R.  Johnson,  A.  M. 
S.  D.  Gromer,  A.  M. 
E.  L.  Morgan,  A.M. 


C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  Ph.  D. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  Sieveking,  B.  S.  in  Agr. 

A.  M.  Cowan,  A.  M. 

AGRICULTURAL  ENGINEERING 

J.  C.  Wooley,  B.  S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

E.  F.  Hopkins,  Ph.  D. 

DAIRY  HUSBANDRY 
A.  C.  Ragsdale,  B.  S.  in  Agr. 

W.  W.  SwETT,  A.  M. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

ENTOMOLOGY 
Leonard  Haseman.  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride,  B.  S.  in  Agr. 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm.  A.  M. 

L.  J.  Stadler,  Ph.  D. 

O.  W.  Letson,  B.  S.  in  Agr. 

Miss  Regina  Schulte* 


Ben  H.  Frame,  B.  S.  in  Agr. 


HORTICULTURE 

V.  R.  Gardner,  M.  S.  A. 

H.  D.  Hooker,  Tr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  Ph.  D. 

F.  C.  Bradford,  M.  S. 

H.  G.  Swartwout,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY 
H.  L.  Kempster,  B.  S. 

Earl  W.  Henderson,  B.S. 

SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M 

W.  A.  Albrecht.  Ph.  D. 

F.  L.  Duley,  A.M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr 
Richard  Bradfield,  Ph.  D. 

VETERINARY  SCIENCE 
J.  W.  Connaway,  D.  V.  S.,  M.  D. 
L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 


OTHER  OFFICERS 

R.  B.  Price,  M.  S..  Treasurer 
Leslie  Cowan,  B.  S..  Secretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B..  Agricultural  Editor 
J.  F.  Barham.  Photographer 

Miss  Jane  Frodsham,  Librarian. 

E.  E.  Brown,  Business  Manager. 


In  service  of  U.  S.  Department  of  Agriculture. 


Relation  of  Temperature  to  Blossoming 
in  the  Apple  and  the  Peach 

F.  C.  Bradford 

Observation  of  the  responsiveness  of  plants  to  certain  tem- 
peratures and  of  the  poleward  progress  of  vegetative  activity  more 
or  less  concurrently  with  the  advance  of  warm  weather  led  to  the 
formulation  many  years  ago  of  the  doctrine  of  thermal  constants. 
According  to  this  theory  a given  stage  in  the  development  of  any 
plant  is  reached  when  that  plant  has  received  a certain  amount  of 
heat,  regardless  of  the  time  required  or  of  the  temperatures  in- 
volved. For  each  plant  and  for  each  successive  stage  there  was 
assumed  to  be  a definite  heat  requirement,  which  generally  re- 
ceived a mathematical  expression  in  the  form  of  so-called  “heat 
units.”  The  unit  was  a degree  on  one  of  the  several  thermometer 
scales.  However  taken,  temperature  observations  were  generally 
reduced  to  terms  of  average  or  mean  daily  temperatures.  The 
readings  for  all  the  days  involved  in  the  period  in  question  were 
combined  and  the  sum  called  the  “thermal  constant,”  since  it  was 
assumed  to  be  constant  for  the  plant  wherever  and  whenever 
grown.  Units  based  on  this  system  may  be  designated  “day-de- 
grees” to  distinguish  them  from  the  “hour-degrees”  obtained  by 
computation  from  hourly  temperatures. 

Enunciated  first,  probably,  by  Reaumer29  in  1735,  the  original 
conception  has  been  modified  by  later  workers.  Adanson2  pointed 
out  that  temperatures  below  freezing  do  not  reverse  plant  activity 
and  discarded  them  from  his  summations.  Others  used  as  bases 
of  calculation  some  still  higher  temperature,  at  which  vegetative 
activity  supposedly  began.  The  number  of  heat  units  for  one  day 
was  obtained  by  subtracting  the  base  temperature  from  the  actual 
— mean  or  maximum  as  the  case  might  be — thermometer  reading 
for  that  day;  consequently  this  has  been  called  the  “remainder 
system.”  Later  graduated  values  were  assigned  to  various  tem- 
peratures in  recognition  of  accelerated  growth  at  certain  tempera- 
tures. The  Livingstons,22  particularly,  have  worked  out  a scale 
of  weighted  temperature  values  based  on  the  principle  of  Van’t  Hoff  and 
Arrhenius,  pointing  out,  however,  that  the  purely  physical  proces- 
ses involved  in  growth  are  not  governed  by  this  principle.  This 
system  they  called  the  “exponential.”  More  recently  Livingston21 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


evolved  a third  system  called  “physiological”,  based  on  Lehen- 
bauer’s  observations  of  root  growth  in  maize  at  various  tempera- 
tures. This  system  differs  from  the  others  in  that  it  recognizes  an 
optimum  temperature  above  which  the  values  assigned  decrease. 
It  is  put  forward,  evidently,  only  as  tentative,  since  Livingston 
states  several  qualifications  of  its  applicability. 

Progress  in  plant  physiology  and  particularly  the  recogni- 
tion that  numerous  factors  influence  plant  growth  have  modified 
the  original  conception  of  thermal  constants.  Numerous  objec- 
tions to  the  original  conception  have  been  stated  aptly  by 
Schimper,31  and  its  rigid  application  is  not  often  attempted,  ex- 
cept in  the  use  of  growing  season  summations  to  characterize  var- 
ious regions,  as  exemplified  by  Merriam’s24  work  on  life  zones, 
and  Swingle’s33  on  the  date  palm.  Ihne,18  though  inclined  to  con- 
sider phenological  observations  a measure  of  the  weather,  express- 
ly repudiates  the  assignment  of  definite  thermal  constants  to  any 
plant. 

Unfortunately  there  has  survived  a supposed  connotation  be- 
tween the  old  thermal  constant  conception  and  phenology  which 
has  retarded  the  study  of  phenological  observations  in  ways  that 
might  otherwise  have  been  attempted.  One  purpose  of  the  present 
paper  is  to  point  out  how  the  thermal  constant  conception,  though 
full  recognition  be  accorded  the  many  objections  to  it,  still  may 
furnish,  in  connection  with  phenological  observations,  a valuable 
tool  in  the  study  of  the  response  by  plants  to  some  of  the  factors 
composing  climate.  The  comparative  meagerness  of  the  available 
data  and  the  limited  number  of  localities  they  represent  preclude 
the  possibility  of  formulating  much  that  is  conclusive,  and  this 
paper  can  be  regarded  only  as  suggestive  of  what  might  be  at- 
tempted with  abundant  data  for  the  same  plant  under  many  con- 
ditions. 

PHENOLOGY  OF  FRUIT  TREES  IN  NORTH  AMERICA 

Systematic  observations  on  the  blossoming  of  fruit  trees  at 
various  points  in  North  America  began,  perhaps,  in  1817  when 
Bigelow7  compiled  a list  of  the  dates  of  blossoming  of  the  peach  at 
various  points  from  Fort  Claiborne,  Alabama,  to  Montreal.  Early 
reports  of  the  Smithsonian  Institution  and  of  the  Army  Signal  Ser- 
vice, the  forerunner  of  the  present  Weather  Bureau,  contain  many 
scattered  observations  on  blossoming  dates.  Following  the  recog- 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  5 


nition  of  the  importance  of  cross  pollination  in  many  fruits,  blos- 
soming data  have  been  published  by  a number  of  agricultural  ex- 
periment stations.  Since  these  were  intended  merely  to  show 
the  overlapping  in  blossoming  seasons  of  horticultural  varieties, 
complementary  temperature  data  are  not  ordinarily  available  and 
study  of  them  shows  little  beyond:  (1)  a general  similarity  in  se- 
quence of  species  and  of  varieties,  (2)  differences  between  places 
in  the  average  lengths  of  the  blossoming  seasons  and  in  the  in- 
tervals between  the  blossoming  of  the  several  fruits,  and  (3)  a 
general,  though  not  uniform,  recession  of  the  blossoming  dates 
with  increased  latitude  and  altitude. 

The  Initial  Date. — Those  who  have  attempted  to  fit  thermal 
constants  to  phenological  observations  on  perennial  plants  have 
found  much  perplexity  in  fixing  an  initial  date  for  temperature 
summations.  Some  have  computed  from  leaf  fall  in  the  previous 
autumn,  some  from  the  coldest  period  of  the  winter  (which  is,  in 
many  cases,  early  in  February)  and  some  from  the  date  when 
the  average  daily  temperature  rises  above  the  freezing  point. 
Others,  as  Fritsch,15  have  considered  the  precise  date  of  little  im- 
portance and  have  used  January  1,  as  a matter  of  convenience. 
This  is,  perhaps,  the  most  commonly  used  starting  point. 

It  is  interesting  that  in  many  plants  heat  summations  from  this 
date  to  blossoming  are  not  the  same  everywhere.  Waugh34  found 
in  1898  a general  tendency  for  blossoming  at  lower  summations 
in  the  north  with  the  “American  wild  plum”,  than  in  the  south. 
More  pronounced  differences  are  evident  in  the  summations  for 
the  Late  Crawford  peach  at  Pomona,  California,  and  at  Wauseon, 
Ohio,  shown  in  Table  1.  These  are  compiled  from  reports  of  the 
California  Agricultural  Experiment  Station9’ 10>  11  and  from  the 
Mikesell  records.25  Though  the  California  figures  are  calculated 
from  monthly  means  of  daily  maximum  temperatures  and  the  Ohio 
figures  from  daily  maximum  readings,  errors  arising  from  this 
cause  must  be  slight  in  proportion;  and  the  great  differences  shown 
in  heat  summations  to  blossoming  actually  exist.  The  highest 
summation  from  January  to  blossoming  for  any  of  the  27  years 
of  the  Ohio  records  is  925,  considerably  below  the  lowest  shown 
here  for  Pomona  and  the  minimum  for  Wauseon  is  barely  more 
than  one-fourth  the  maximum  for  Pomona. 


6 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


Table  1. — Heat  Summations  in  Day-Degrees  (Maximum  Above  43)  eor  the 
Tate  Crawford  Peach  From  January  1 to  Blossoming  in  Ohio 
and  in  California. 


Year 

Wauseon,  Ohio 

Pomona,  California 

Date  of  first 
blossoming 

Day- 

degrees 

Date  of  first 
blossoming 

Day- 

degrees 

1894 

Apr.  17 

745 

Mar.  15 

1222 

1895 

May  2 

860 

Mar.  2 

1266 

1896 

Apr.  23 

650 

Mar.  20 

2217 

1902 

Apr.  29 

804 

Apr.  1 

2329 

1903 

Mar.  25 

1895 

Average 

(27  yrs.) 

732 

(5  yrs.) 

1786 

There  may  be,  then,  a considerable  and  consistent  inequality 
in  heat  summations  from  January  1 to  blossoming  for  the  same 
fruit  grown  at  points  differing  considerably  in  climate.  This  is 
in  accord  with  observations  of  Palladin,27  who  records  similar  dif- 
ferences in  many  plants  at  Brussels  and  at  Petrograd ; these  dif- 
ferences were  much  more  pronounced  in  the  early  blossoming 
than  in  the  late  blossoming  plants.  According  to  one  view,  ad- 
vanced by  Linsser20  the  total  heat  requirements  for  any  stage 
of  plant  development  are  not  identical  at  all  places,  but  their 
proportion  to  the  total  heat  summation  of  the  year  is  everywhere 
constant.  In  other  words,  there  is  supposed  to  be  an  acclimatiza- 
tion so  that  the  same  function  may  be  performed  with  less  heat 
at  one  point  than  at  another,  but  require  the  same  proportion  to 
the  total  for  the  year  at  all  points.  This  hypothesis  appears  un- 
tenable, in  some  cases  at  least,  since  this  numerical  ratio  between 
the  accumulation  to  ripening  in  the  peach  and  the  total  for  the 
year  varies  widely,  from  48.8  per  cent  in  Alabama  to  83.1  in 
Massachusetts.16  Furthermore,  it  does  not  take  into  account  sea- 
sonal variations  at  the  same  point.  Seeley32  found  great  fluctua- 
tions from  year  to  year  in  heat  accumulations  for  various  epochs 
in  the  Late  Crawford  peach  at  Wauseon,  Ohio.  In  some  years 
the  minimum  accumulation  was  70  per  cent  of  the  maximum  for 
the  same  period,  in  another  50  and  in  one,  only  38.  Evidently, 
then,  Linsser’s  constant  or  “aliquot”  will  not  explain  such  differ- 
ences as  those  in  summations  in  California  and  in  Ohio  from  Janu- 
ary 1 to  blossoming.  Finally,  since  heat  accumulations  at  one 
point  may  vary  considerably  from  year  to  year,  if  carried  to  ex- 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  7 


tremes  this  hypothesis  implies  a rather  remarkable  prescience  in 
the  plant. 

The  differences  between  localities  in  heat  accumulations  to 
blossoming  may  be  due  in  part  to  different  normal  temperature 
distributions.  Price28  demonstrated  an  acceleration  in  blossom- 
ing of  peach  and  plum  with  high  temperatures;  yet  his  data  show 
that  the  twigs  held  at  the  lower  temperatures,  though  they  re- 
quired a longer  time  for  blossoming,  actually  received  in  some 
cases  less  total  heat  (in  day-degrees).  In  some  localities  it  is 
possible  that  even  before  blossoming  there  occur  temperatures  high 
enough  to  exercise  an  inhibitory  effect,  or,  perhaps,  the  winters 
are  not  cold  enough  to  make  subsequent  high  temperatures  fully 
effective.  Twigs  of  ash  and  linden  cut  before  the  end  of  the  rest 
period  were  kept  by  Weber35  in  a dormant  state  in  a warm  green- 
house for  15  months;  at  the  end  of  this  time  most  of  the  buds 
opened  normally. 

VARIABILITY  IN  SUMMATIONS  TO  BLOSSOMING 

Since  fruit  bud  differentiation  in  several  fruits  is  first  evident 
about  July  1,  it  has  been  suggested  that  summations  should  be  computed 
from  this  time  to  blossoming  in  the  following  spring.  If  this  date  is 
used  for  beginning  computations  on  the  apple  and  on  the  plum  in 
Wisconsin,  there  is  apparently  a closer  agreement  from  year  to 
year  than  when  summations  are  made  from  January  1 ; this  has 
been  interpreted  to  indicate  July  1 as  the  proper  starting  point.30 
Much  of  this  apparent  agreement,  however,  is  due  to  the  tendency 
of  meteorological  elements  to  average  alike  over  long  periods. 
Summations  calculated  from  July  1 to  the  following  May  1,  the 
approximate  date  of  fruit  bud  opening,  fit  very  nearly  as  closely 
as  those  figured  to  the  dates  of  actual  blossoming.  The  ratio  be- 
tween the  smallest  summation  and  the  largest  is,  in  the  Doney 
plum,  86.8  per  cent;  in  the  calendar  summations,  the  check,  it  is 
82.9  per  cent. 

The  very  fact  that  the  Doney  plum  came  into  blossom  in  1904 
with  4,494  day-degrees  from  July  1 while  in  1901  the  accumulation 
was  5,174,  or  680  more,  suggests  that  in  the  latter  year  some  heat 
was  received  when  it  was  ineffective  in  forwarding  blossoms  or 
was  received  in  surplus  quantities  or  that  at  some  periods  in  the 
cycle  heat  is  not  a controlling  factor. 

Indeed,  something  of  the  sort  may  be  deduced  from  the  data 
published  by  Sandsten.  If  the  summations  from  successive  dates 


8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


be  averaged  and  their  respective  mean  deviations  determined,  it 
becomes  evident  that  the  ratio  betwen  the  average  of  the  sum- 
mations and  the  mean  deviation  (in  other  words,  the  variability 
of  the  summations)  changes  and  that  it  does  not  diminish  in  strict 
accordance  with  the  tendency  of  meteorological  values  toward 
greater  uniformity  with  increasing  time.  This  is  shown  in  Table 
2,  arranged  from  Sandsten’s  data  for  the  Forest  Garden  plum, 
where  the  ratio  just  mentioned  is  designated  the  coefficient  of 
variability.  The  “coefficient  of  variability”  used  in  this  paper  is 
calculated  from  the  mean,  rather  than  from  the  standard,  devia- 
tion, to  lessen  the  effects  of  extreme  variations.37 


Table  2. — Heat  Summations  in  the  Forest  Garden  Plum  at  Madison.  Wis- 
consin, 1900-1905  Inclusive. 

(Compiled  from  data  by  Sandsten30) 


To  blossoming 
from 

Mean  of 
summations 

Average 

deviation 

Coefficient  of 
variability 

July  1. 

4836 

181 

3.74 

Aug.  1 

3608 

157 

4.35 

Sept.  1 

2416 

71 

2.93 

Oct.  1 

1540 

97 

6.29 

Nov.  1 

893 

80 

8.96 

Dec.  1 

678 

61 

9.99 

Jan.  1 

667 

70 

10.49 

Feb.  1 

662 

71 

10.72 

Mar.  1 

653 

63 

9.64 

Apr.  1 

523 

58 

11.08 

Sept.  1 (omitting 

Nov., 

Dec.,  Jan.,  and 

Feb.)  2159 

50 

2.31 

The  significance  of  these  coefficients  is  more  apparent  if  they 
are  studied  beginning  with  the  coefficient  for  January  1.  On 
either  side  of  this  (December  and  March)  are  lower  values,  signi- 
fying  greater  agreement  in  summations  beginning  at  other  times 
and  suggesting  that  temperature  accumulations  from  this  date  are 
not  altogether  effective  in  advancing  blossoming.  In  summations 
beginning  March  1 there  is  closer  agreement;  and  the  high  varia- 
bility from  April  1 may  be  interpreted  to  mean  that  advancement 
toward  blossoming  has  begun,  in  some  years  at  least,  by  that  time. 
The  difference  in  coefficients  between  November  1 and  October 
1 is  striking  and  suggests  that,  beginning  possibly  about  November 
1,  the  temperatures  received  are  not  ordinarily  effective.  The 
greatest  agreement  in  summations  in  the  whole  series  is  in  those 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  9 


dating  from  September  1 ; the  low  coefficient  at  this  point  is  re- 
markable. If,  however,  the  November,  December,  January  and 
February  temperatures  are  omitted  the  coefficient  of  variability  in 
summations  from  this  date  is  diminished  even  further. 

From  August  1 and  July  1,  though  these  months  in  them- 
selves usually  show  relatively  slight  variability  in  their  tempera- 
ture summations,  the  coefficients  of  variability  are  higher.  Their 
low  value  as  compared  with  that  of  March  is  due  to  the  longer 
period  covered,  and  their  significance  is  probably  slight. 

Here,  then,  though  caution  must  be  observed  against  infer- 
ring too  much,  there  seems  to  be  reason  to  consider  tentatively  for 
this  fruit  at  Madison:  (1)  that  temperature  deficiency  during 

July  and  August  is  not  a limiting  factor  in  any  ordinary  season, 
(2)  that  it  becomes  in  some  measure  a limiting  factor  during  Sep- 
tember and  October,  (3)  that  temperature  is  ineffective  during 
November,  December,  January  and  February,  possibly  because 
there  is  not  enough  heat  received  to  have  any  appreciable  effect 
and  (4)  that  about  March  1 it  again  becomes  for  a time  a determ- 
ining factor.  Under  other  conditions,  of  course,  very  high  temper- 
atures may  become  limiting. 

Even  though  these  indications  be  true  for  the  Forest  Garden 
plum,  caution  should  be  exercised  in  applying  them  to  another 
plant,  for  example  a Japanese  plum,  in  Wisconsin,  or  to  the  same 
plum  in  another  locality.  In  other  words,  significant  dates  for 
phenological  data  may  conceivably  differ  with  the  plant  and  with 
the  locality.  Angot3  carried  this  idea  of  flexibility  to  the  extreme, 
stating  that  the  significant  date  varied  not  alone  with  the  plant 
and  the  locality  but  also  from  year  to  year.  Evidence  is  intro- 
duced in  this  paper  indicating  the  variation  of  the  significant  date 
with  plant  and  with  locality;  as  to  the  yearly  variation  in  the 
same  plant  and  in  the  same  locality  the  evidence  is  less  clear.  If, 
however,  the  chemical  composition  of  the  plant  be  considered  to 
have  an  influence,  as  seems  quite  plausible,  the  effective  date  may 
vary  as  well.  Furthermore,  the  stage  of  blossom  development  at- 
tained in  the  fall  has  been  shown  by  Magness23  to  vary  from  year 
to  year  in  the  same  variety.  Consequently  some  variation  in 
opening  in  the  spring  might  be  expected  even  in  seasons  that  pre- 
sent practically  the  same  temperatures. 

THE  APPLE  AND  THE  PEACH  IN  OHIO 

A study  covering  a number  of  seasons  at  one  point  has  cer- 


10 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


tain  advantages  over  studies  of  a few  seasons  at  many  points.  If, 
for  example,  the  date  when  temperatures  become  effective  be 
conceived  to  vary  from  place  to  place  there  is  no  satisfactory  way 
of  ascertaining  this  date  from  scattered  observations  unless  the 
minimum  accumulation  to  blossoming  observed  at  any  point  be 
subtracted  from  the  accumulations  at  other  points  and  the  dates 
computed  from  the  day-degree  remainders.  This  is,  in  effect,  shap- 
ing the  problem  to  fit  the  answer.  In  observations  at  one  point 
over  a series  of  years  a certain  degree  of  variation  in  other  limit- 
ing factors  is  presumably  reduced  and  if  there  is  any  validity  in 
the  thermal  constant  conception  it  should  appear  in  observations 
of  this  sort. 

The  publication  by  the  United  States  Weather  Bureau  of  the 
Mikesell  records,25  comprising  phenological  observations  on  num- 
erous plants  at  Wauseon,  Ohio,  over  a period  of  30  years,  to- 
gether with  daily  meteorological  records,  makes  possible  a rather 
critical  comparison  of  heat  accumulations  and  phenological  obser- 
vations. 

Since  these  records  cover  a longer  period  than  any  other  avail- 
able data  they  are  analyzed  here  and  used  in  the  study  of  the 
records  of  the  Missouri  Agricultural  Experiment  Station,  which 
cover  a much  shorter  time. 

Methods  Used. — Heat  accumulations  may  be  measured  in  var- 
ious ways.  In  the  work  reported  here  the  simple  summations  of 
temperature  to  blossoming,  both  maximum  and  mean,  above  sev- 
eral thermometric  points,  were  computed.  In  addition  one  series 
was  computed  on  the  exponential  system.  For  each  series  the 
yearly  summations  were  averaged,  the  mean  deviations  from  the 
averages  determined  and  variability  coefficients  derived  by  divid- 
ing the  mean  deviations  by  the  averages  of  the  total  accumula- 
tions. Occasional  trials  showed  no  material  relative  changes  in 
coefficients  resulting  from  the  use  of  standard  or  mean  deviations. 
The  coefficients  derived  by  the  several  methods  are  shown  in 


Table  3. — Variability  Coefficients  of  Heat  Summations  From  January  1 
to  Blossoming  at  Wauseon,  Ohio,  as  Calculated  on  Different  Bases. 


Base 

System 

Apple 

Peach 

32°F.  Max. 

Remainder 

7.69 

8.26 

43 °F.  Max. 

Remainder 

8.79 

9.80 

50°F.  Max. 

Remainder 

10.48 

12.73 

43 °F.  Mean 

Remainder 

12.20 

16.06 

40 °F.  Max. 

Exponential 

10.38 

9.97 

Relation  of  Temperature  to  Blossoming — Apple  and  Peach  11 


The  magnitude  of  the  variability  seems  to  vary  inversely  with 
the  number  of  units  involved ; for  this  reason  the  lower  variabil- 
ity resulting  from  the  use  of  32°  as  the  base  point  is  not  neces- 
sarily significant.  Since  this  comparison  did  not  show  any  base- 
point  or  system  to  be  markedly  superior  to  any  other,  the  series 
based  on  maximum  temperature  above  43°  was  chosen  for  most  of 
the  further  computations.  This  system  appeared  to  give  inter- 
mediate values  and  its  results  would  be  comparable  with  other 
work  which  has  been  based  on  the  same  temperature. 

Temperature  observations  taken  according  to  conventional 
meteorological  methods  are  not  true  records  of  plant  tempera- 
tures and  since  the  disparity  between  the  two  varies  no  correc- 
tions can  be  applied.  For  present  purposes,  however,  since  in 
sunshine  twigs  are  generally  warmer  than  the  air,  maximum  air 
temperatures  probably  approximate  those  of  the  plant  more  closely 
than  mean  air  temperatures.  For  other  seasons  or  for  other  tem- 
perature ranges  or  in  other  climates  mean  temperatures  might  be 
preferable.  However,  even  on  summer  stages  for  the  peach  at 
Wauseon,  Seeley32  found  less  variation  in  computations  involving 
maximum  than  in  those  involving  mean  temperatures,  though  it 
is  true  this  lower  figure  may  be  due  to  the  larger  number  of  units 
involved. 

Calculations. — Though  it  seems  unlikely  that  heat  deficiency  is 
a limiting  factor  with  apples  or  peaches  in  Ohio  during  the  sum- 
mer months,  computations  were  made  from  July  1 to  blossoming 
the  following  year.  From  these  figures  the  summations  from 
other  dates  to  blossoming  were  readily  secured  and  the  respec- 
tive variability  coefficients  determined.  As  a check  on  these,  sum- 
mations to  April  28  and  to  May  7,  the  average  blossoming  dates 
of  the  peach  and  of  the  apple,  respectively,  were  similarly  com- 
puted. These  may  be  considered  as  measures  of  the  independent 
variability  of  the  weather  and  are  valuable  for  comparison  with 
the  variability  to  the  actual  dates  of  blossoming. 

If  heat  accumulations  are  plotted  vertically  and  a horizontal 
scale  be  adopted  for  time  such  that  the  spread  of  the  projections 
of  blossoming  dates  on  the  abscissa  is  equal  in  length  to  the  spread 
of  the  projections  of  the  accumulations  on  the  ordinate,  mathe- 
matical expression  of  the  trend  of  the  line  connecting  blossoming 
dates  is  possible.  This  is,  in  effect,  done  when  the  coefficient  of 
variability  in  accumulations  to  blossoming  is  divided  by  the  coef- 
ficient of  variability  to  the  average  date  of  blossoming.  With  per- 


12 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


feet  uniformity  in  total  day-degrees  to  blossoming  the  line  would 
be  horizontal ; with  perfect  uniformity  in  totals  to  a given  date 
the  line  would  be  vertical.  With  an  equal  degree  of  uniformity 
in  both  it  would  be  at  a slope  of  45°.  This  would  be  the  case 
were  coefficients  of  variability  to  blossoming  and  to  average  date 
of  blossoming  equal. 

In  short,  the  numerical  ratio  obtained  by  dividing  the  coef- 
ficient of  variability  in  summations  to  blossoming  by  the  coef- 
ficient of  variability  in  summations  to  average  date  is  the  tan- 
gent of  the  angle  with  the  horizontal  made  by  a smooth  line  con- 
necting the  dates  of  blossoming.  When  this  ratio  is  above  one, 
the  angle  is  greater  than  45°  and  nearer  vertical.  In  other  words, 
the  agreement  is  closer  with  the  average  date  than  in  the  total 
accumulations. 

Accordingly  the  figures  in  the  columns  headed  “Tangent” 
in  Table  4 are  in  reality  tangents  of  slopes  of  lines  connecting 
the  graphical  positions  of  the  blossoming  dates.  The  value  0.94 
for  the  apple  indicates  a slope  of  approximately  43°  for  this  line- 
practical  neutrality.  The  value  0.51  indicates  a slope  of  ap- 
proximately 27.°  In  the  peach  the  1.47  value  indicates  a slope  of 
56°,  nearer  vertical  than  horizontal.  However,  even  without  ex- 
pression in  degrees,  the  tangents  serve  for  comparison. 


Table  4. — Variability  Coefficients  of  Heat  Accumulations  From  Various 
Dates  to  Blossoming  in  the  Apple  and  in  the  Peach  at  Wauseon,  Ohio. 


Beginning 

date 

Apple 

Peach 

To  actual 
blossom- 
ing 

To  average 
date  of 
blossoming 

Tangent 

To  actual 
blossom- 
ing 

To  average 
date  of 
blossoming 

Tangent 

July  1 

4.55 

5.36 

0.85 

5.27 

5.00 

1.05 

Aug.  1 

5.63 

6.45 

0.87 

6.22 

6.34 

0.98 

Sept.  1 

7.37 

7.86 

0.94 

8.04 

7.66 

1.05 

Oct.  1 

9.33 

11.11 

0.84 

9.71 

10.59 

0.92 

Nov.  1 

11.08 

13.80 

0.80 

7.77 

14.33 

0.54 

Dec.  1 

13.65 

15.47 

0.88 

9.55 

16.94 

0.56 

Jan.  1 

8.79 

15.64 

0.56 

9.80 

16.37 

0.60 

Feb.  1 

8.48 

15.78 

0.54 

11.34 

16.73 

0.68 

Mar.  1 

9.79 

16.76 

0.58 

13.40 

17.51 

0.76 

Mar.  15 

7.93 

15.51 

0.51 

14.91 

15.60 

0.95 

Apr.  1 

14.70 

15.79 

0.93 

25.22 

17.09 

1.47 

Relation  of  Temperature  to  Blossoming — Apple  and  Peach  13 


Indications. — The  lower  the  tangent  the  more  significant,  pre- 
sumably, is  the  coefficient  for  the  corresponding  period.  Conse- 
quently the  low  variability  coefficients  for  the  summer  and  au- 
tumn months  lose  their  weight  and  those  of  some  of  the  later 
dates  become  more  significant. 

An  interesting  difference  between  the  apple  and  the  peach  is 
revealed  by  inspection  of  the  tangents.  The  lowest  value  in  the 
peach  is  in  the  summations  figured  from  November  1 ; in  the  apple 
the  lowest  value  is  in  the  figures  dating  from  March  15.  This  dif- 
ference seems  to  indicate  that,  under  the  conditions  obtaining  at 
Wauseon,  high  temperatures  during  winter  are  effective  in  promot- 
ing growth  in  the  peach,  but  not  in  the  apple. 

In  the  apple  the  period  of  effective  temperatures  seems  more 
definitely  fixed  than  in  the  peach.  From  January  1 to  March  15 
in  the  apple  the  tangents  change  but  little,  with  the  smallest  fig- 
ure on  March  15.  It  should  be  considered,  however,  that  heat 
accumulations  are  small  during  this  time  and  can  affect  the  total 
variability  but  little.  Other  things  equal,  such  changes  as  do 
occur  as  the  date  of  summations  moves  backward  should,  through 
augmenting  somewhat  the  total  of  day-degrees  involved,  reduce 
the  variability.  Therefore  even  the  slight  difference  in  tangents 
shown  may  be  significant  in  the  apple.  The  occurrence  of  the 
lowest  figure  on  March  14  does  not  signify  that  the  rest  period 
ends  then.  It  is,  in  a sense,  an  average  date  and  means  that, 
broadly  speaking,  advancement  starts  in  half  the  years  at  that 
time.  Consequently  the  end  of  the  rest  period  must  be  earlier. 

In  the  peach,  the  succession  of  low  values  is  in  the  reverse 
order  and  so  far  as  this  array  affords  evidence,  the  decrease  from 
January  1 or  December  1 may  be  due  merely  to  the  longer  period 
and  the  consequently  greater  total  of  units  involved.  In  either 
case,  however,  it  seems  clear  that  the  peach  becomes  responsive 
to  high  temperature  earlier  than  the  apple  and  that  its  earlier 
blossoming  is  not  necessarily  due  to  a lower  total  requirement 
of  heat.  The  low  temperature  of  the  ordinary  winter  at  Wauseon 
would  keep  the  trees  dormant  and  microscopic  study  of  buds  for 
several  years  might  show  no  development  during  this  time,  unless 
the  period  of  investigation  happened  to  include  a mild  winter. 
Dormancy  of  the  peach  in  the  north  and  in  the  south  may  be 
quite  different;  in  the  one  case  imposed  by  low  temperature  and 
in  the  other  by  the  rest  period.  Johnston19  found  that  the  mois- 
ture content  of  peach  buds  in  Maryland  increases  after  January  1 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


in  a definite  relationship  to  the  “sum  of  the  effective  daily  mean 
temperature  above  43°. ” 

Seasonal  Differences. — Illustration  of  the  difference  between 
the  two  types  of  fruit  is  found  in  the  graphs  of  heat  accumula- 
tions from  January  1 in  1890  and  in  1912,  shown  in  figure  1.  These 
years  are  selected  because  they  represent  respectively  the  maxi- 
mum and  the  minimum  accumulations  of  heat  from  January  1 to 
March  1.  In  1912,  with  little  accumulation  of  heat  prior  to  April 
1,  the  apple  came  into  blossom  very  close  to  the  peach  both  in 
time  and  in  heat  accumulations.  This  year,  in  fact,  marked  the 


Fig.  1. — Blossoming  of  peach  and  apple  in  years  of  maximum  and  of  minimum 
accumulation  on  March  1,  at  Wauseon,  O. 


lowest  summation  to  blossoming  for  the  apple.  In  1890,  with  con- 
siderable heat  accumulation  throughout  the  winter  months,  the 
peach  came  into  blossom  earlier,  but  with  substantially  the 
same  heat  accumulation  as  in  1912.  The  apple,  however,  though 
its  date  of  blossoming  was  nearly  the  same  as  in  1912,  had  re- 
ceived 300  day-degrees  more  of  heat.  This  difference  is  about 
the  same  as  the  margin  by  which  the  accumulation  to  April  1 
in  1890  exceeded  that  of  1912.  This  may  be  interpreted  to  signify 
that  in  1912  practically  all  the  heat  received  came  when  it  was 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  15 


effective,  while  in  1890  much  of  it  was  ineffective  for  the  apple. 
Apparently  the  peach  started  from  dormancy  earlier  than  the  apple 
in  1890,  while  in  1912  there  was  little  difference,  because  the  low 
temperatures  held  both  dormant. 

Since  these  two  winters  were  so  widely  different,  it  seems 
logical  to  infer  that  localities  with  average  winters  differing  (as 
do  these  extreme  types)  would  show  for  regions  with  mild  winters 
a considerable  spread  in  blossoming  season  between  the  peach 
and  the  apple — well  known  to  be  the  case — while  those  with  cold 
winters  would  show  little  or  no  difference.  In  extreme  cases  the 
peach  and  the  apple  may  bloom  simultaneously.  This  condition 
occurred  at  Wauseon  in  1895,  following  a cold  January  and  Feb- 
ruary and  was  closely  approached  in  other  years,  invariably  fol- 
lowing winters  of  small  heat  accumulations.  Indeed,  in  1912 
which,  according  to  Hedrick17  was  not  an  unusual  blossoming  sea- 
son, at  Geneva,  New  York,  blossoming  in  the  apples  began  a 
day  ahead  of  the  peaches.  The  same  phenomenon  occurred  in 
1905  at  Columbia,  Missouri.  The  variations  sometimes  reported 
in  the  sequence  of  blossoming  in  other  fruits  may  be  due  to  simi- 
lar causes. 

If  a certain  validity  be  assumed  for  the  thermal  constant  con- 
ception, the  rather  wide  difference  from  year  to  year  in  the  sum- 
mations from  any  given  date  to  blossoming  suggest  that  the 
higher  figures  may  be  due  to  accumulations  occurring  during  per- 
iods when  they  are  ineffective  or  in  greater  quantities  than  can  be 
fully  effective — and  of  course  other  factors  than  temperature  may 
intervene.  To  facilitate  comparison,  data  for  the  years  of  maxi- 
mum and  of  minimum  accumulations  from  January  1 to  blossom- 
ing in  the  peach  and  in  the  apple  are  arranged  in  Table  5.  It  is 
interesting  and  significant  that  these  years  are  not  identical  for 
the  two  fruits,  only  three  duplications  occurring.  In  both  fruits 
the  minimum  accumulations  from  January  1 to  blossoming  average 
practically  the  same  in  relation  to  the  maximum,  but  here  the  sim- 
ilarity stops.  In  the  peach  the  years  of  lowest  summation  from 
January  1 to  blossoming  generally  succeed  periods  of  consider- 
able accumulation  in  November  and  December,  the  accumulations 
preceding  the  years  of  minimum  accumulation  being  in  fact  141 
per  cent  of  those  preceding  the  maximum.  In  the  apple  it  is  ap- 
parently a matter  of  indifference,  since  the  November  and  De- 
cember accumulations  preceding  the  minimum  and  the  maximum 
years  average  practically  the  same.  It  should  be  stated  that  the 


16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


Table  5. — Analysis  oe  Accumulations  in  Years  oe  Greatest  and  oe  Lowest 
Summations  From  January  1 to  Blossoming  in  the  Peach  and  in 
the  Apple  at  Wauseon,  Ohio. 

(In  day-degrees) 


Year 

Jan.  1 to 
blossoming 

Previous  Nov. 
1 to  Jan.  1 

Jan.  1 to 
Mar.  1 

Jan.  1 to 
Mar.  15 

Mar.  15  to 
blossoming: 

Peach 

Years  of  Minimum  Summations 

1910 

632 

363 

14 

112 

520 

1908 

588 

180 

18 

104 

484 

1900 

627 

305 

86 

100 

527 

1896 

650 

286 

50 

57 

593 

1891 

594 

296 

125 

142 

452 

Years  of  Maximum  Summations 

1902 

804 

184 

52 

144 

660 

1898 

834 

266 

72 

198 

636 

1895 

860 

222 

31 

45 

815 

1893 

853 

150 

13 

104 

749 

1887 

870 

195 

80 

213 

657 

Apple 

Years  of  Minimum  Summations 

1912 

752 

195 

6 

6 

746 

1908 

791 

302 

18 

104 

687 

1905 

787 

290 

13 

15 

772 

1896 

779 

286 

12 

57 

722 

1886 

803 

217 

83 

196 

607 

Years  of  Maximum  Summations 

1901 

1005 

259 

42 

66 

939 

1894 

1217 

323 

124 

324 

893 

1890 

1052 

296 

294 

327 

725 

1889 

1056 

308 

70 

148 

908 

1887 

1079 

195 

80 

213 

866 

Averages 

Peach 

Min. 

618 

286 

59 

103 

515 

Max. 

844 

203 

49 

141 

703 

Apple 

Min. 

781 

258 

26 

56 

707 

Max. 

1081 

276 

122 

215 

866 

Minimum  in  per  cent  of  Maximum. 

Peach 

73.2 

141 

120 

73 

73 

Apple 

72.2 

93 

21.0 

26 

82 

Relation  of  Temperature  to  Blossoming — Apple  and  Peach  17 


two  years  of  greatest  heat  accumulation  in  November  and  Decem- 
ber were  followed  by  crop  failures  in  the  peach,  constituting  two 
out  of  the  three  in  the  30  years  of  record.  These  facts,  together 
with  the  low  variability  coefficient  in  the  peach  summations  from 
November  1 (Table  4)  indicate  that  rather  marked  accumulations 
of  heat  in  November  and  December  have  some  influence  in  the 
forwarding  of  Late  Crawford  peach  blossoms  toward  opening,  but 
are  not  important  in  the  King  apple. 

Johnston19  found  that  the  relation  between  temperature  ac- 
cumulations from  January  1 and  moisture  content  of  peach  fruit 
buds,  though  constant  in  any  one  year,  varies  from  year  to  year; 
and  that  “certain  conditioning  influences  that  are  operative  dur- 
ing or  preceding  dormancy  apparently  ‘predetermine’  the  exact  re- 
lationship between  air  temperature  and  the  moisture  content  of 
the  buds  for  the  period  following  dormancy.” 

The  averages  in  Table  5 show  an  excess  of  heat  during  Jan- 
uary and  February  of  the  years  of  minimum  accumulation  for  the 
peach,  but  inspection  of  the  detailed  figures  shows  that  this  is  of 
doubtful  significance,  since  it  is  due  to  a high  value  in  one  year 
only.  The  low  ratio  of  the  minimum  to  the  maximum  years  (21 
per  cent)  in  the  apple,  however,  apparently  signifies  that  the 
apple  is  unresponsive  at  this  time  and  that  heat  accumulations 
during  this  period  merely  swell  the  total  without  having  any 
marked  effect  in  advancing  the  blossoms.  The  same  negative  re- 
lationship appears  in  the  figures  to  March  15  for  the  apple  (16 
per  cent)  while  the  figures  for  the  peach  change  markedly  and 
assume  the  same  relationship  as  the  total  accumulations.  The 
similarity  in  the  relationship  in  the  peach  of  the  years  of  maximum 
to  those  of  minimum  accumulations  from  January  1 to  blossom- 
ing, from  January  1 to  March  15  and  from  March  15  to  blossom- 
ing suggests  that  the  same  influences  are  operative  during  all  three 
periods;  in  other  words,  that  development  is  progressing.  In  the 
apple  the  change  at  this  time  is  abrupt — from  28  to  86  per  cent — 
the  accumulations  from  March  15  to  blossoming  being  more  nearly 
alike  as  between  maximum  and  minimum  years  than  those  from 
January  1 and  closer  than  in  the  peach — 86  as  compared  to  73 
per  cent. 

Assuming,  for  the  reasons  given  above,  November  1 to  mark 
the  commencement  of  possible  effective  temperatures  in  advanc- 
ing the  peach  toward  blossoming  and  March  15  for  the  apple, 
data  for  the  five  years  of  maximum  summations  for  these  respec- 


18 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


tive  periods  in  each  fruit  are  assembled  in  Table  6 to  show  their 
relation  to  the  temperatures  of  October,  September  and  August 
preceding. 


Tabus  6. — Temperature  Summations  From  Date  oe  PossibeE  Eeeectiveness 
in  Relation  to  Temperature  oe  Previous  Months,  at  Wauseon,  Ohio. 


Peach 

Apple 

Year 

Nov.  1 
to  blos- 
soming 

Oct. 

Sept. 

Aug. 

Year 

Mar.  15 
to  blos- 
soming 

Oct. 

Sept. 

Aug. 

Years  of  Minimum  Accumulation 


1911-12 

879 

554 

979 

1203 

1910-11 

669 

647 

979 

1245 

1910-11 

782 

647 

979 

1245 

1908-09 

694 

699 

1202 

1249 

1907-08 

768 

380 

932 

1190 

1907-08 

687 

380 

932 

1190 

1906-07 

878 

326 

1154 

1326 

1901-02 

706 

740 

1063 

1350 

1890-91 

890 

502 

839 

1213 

1885-86 

707 

430 

952 

1075 

Years  of  Maximum  Accumulation. 

1897-98 

1100 

882 

1270 

1211 

1900-01 

939 

940 

1174 

1448 

1894-95 

1082 

620 

1109 

1361 

1893-94 

893 

597 

1048 

1317 

1893-94 

1068 

597 

1048 

1317 

1888-89 

908 

476 

958 

1289 

1891-92 

1071 

587 

1201 

1254 

1887-88 

903 

460 

990 

1284 

1883-84 

1104 

378 

866 

1137 

1883-84 

892 

378 

866 

1137 

Averages 

Min.  yrs. 

839 

481 

977 

1235 

693 

579 

1026 

1222 

Max.  yrs. 

1085 

613 

1099 

1256 

907 

550 

1072 

1295 

Minimum 

in  per 

cent  of  Maximum. 

Peach 

77 

78 

89 

98 

76 

105 

96 

94 

These  figures  suggest,  though  not  very  strongly,  a tendency 
toward  an  association  between  lower  temperatures  in  October  and 
a low  summation  from  November  1 to  blossoming  in  the  peach.  In 
the  apple  there  is  little  or  no  appearance  of  any  relationship.  This 
difference  may  possibly  be  associated  with  some  effect  of  the  high 
October  temperatures  in  prolonging  or  of  the  low  temperatures  in 
breaking  the  rest  period  in  the  peach,  while  in  the  apple  at  this 
time  they  have,  ordinarily,  no  apparent  effect.  However,  since 
rainfall  in  September  is  likely  to  be  important  in  connection  with 
September  and  October  temperatures,  no  clear  evidence  is  afforded 
by  the  data  in  Table  6 as  to  the  effects  of  October  temperature, 
though  the  essential  similarity  in  September  and  August  summa- 
tions indicates  that  temperature  variations  in  these  months  have 
little  effect  on  these  fruits  in  this  locality. 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  19 


MISSOURI  RECORDS 

Rather  complete  phenological  records  of  numerous  varieties 
of  apples  and  peaches  were  kept  at  the  Missouri  Agricultural  Ex- 
periment Station  from  1905  to  1918  inclusive,  with  the  exception 
of  the  blossoming  records  for  1910.  This  was  an  early  season 
and  the  records  show  most  varieties  in  full  bloom  on  March  28 
but  the  dates  of  first  blossoming  are  not  recorded ; consequently 
this  year  is  omitted  from  calculations  reported  here. 

Through  the  kindness  of  Mr.  George  Reeder,  of  the  United 
States  Weather  Bureau,  temperature  records  for  the  period  cov- 
ered by  the  phenological  data  have  been  made  available.  These 
observations  were  made  at  the  Weather  Bureau  office,  about  one- 
fourth  mile  from  the  University  Orchard  in  which  the  phenologi- 
cal observations  were  taken. 

An  interesting  commentary  on  the  hazards  of  peach  growing 
in  this  section  is  the  appearance  of  blossoming  dates  for  peaches 
for  only  8 of  the  13  years  of  the  record.  Since  the  observations 
were  made  with  considerable  care  it  is  safe  to  presume  that  no 
blossoms  appeared  in  other  seasons  during  this  period.  Com- 
pared with  the  27  crops  in  30  years  at  Wauseon,  Ohio,  and  with 
the  uninterrupted,  though  brief,  sequence  reported  from  Pomona, 
California,  they  suggest  that  this  particular  section  may  be  termed 
a no-man’s  land  for  the  common  varieties  of  peach,  being  sub- 
jected to  the  hazards  of  both  northern  and  southern  types  of 
winter  injury,  (extreme  cold  and  untimely  warm  weather  respec- 
tively) while  regions  north  and  south  are  subject  ordinarily  to 
only  one  form.  Because  of  the  scarcity  of  data  no  attempt  is 
made  here  to  study  extensively  the  climatic  relations  of  the  peach 
in  central  Missouri. 

The  comparative  brevity  of  the  period  for  which  data  are 
available  at  Columbia  increases  the  difficulty  of  formulating  any 
hypothesis  as  to  the  periods  of  effective  temperatures.  Similarly, 
though  data  are  available  for  a considerable  number  of  varieties, 
the  brevity  of  the  record  for  each  makes  varietal  comparisons  rather 
uncertain.  However,  some  generalizations  seem  safe.  The  warmer 
winter  months  at  Columbia  make  the  average  heat  summations  up 
to  the  date  of  blossoming  greater  than  those  at  Wauseon,  though 
the  difference  is  not  so  marked  as  that  between  California  and 
Ohio  for  the  peach.  Since  the  comparison  in  Table  7 between 
summations  at  blossoming  at  Columbia  and  at  Wauseon  is  be-, 
tween  the  King  apple  at  Wauseon  and  the  Fameuse  at  Colum- 


20 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


Table  7. — Average  Temperature  Accumulations  (Max.  Above  43°)  From 
January  1 to  Blossoming  in  the  Apple  at  Wauseon,  Ohio,  and 
at  Columbia,  Mo. 

Wauseon,  Ohio  Columbia,  Missouri 


To  King  Apple  Fameuse  Apple 


February  1 36  122 

March  1 72  261 

March  15  127  389 

April  1 254  646 

Blossoming  912  950 


bia,  the  actual  difference  in  any  one  variety  would  be  somewhat 
greater.  It  is  interesting  that  Fameuse  blossomed  in  1895,  ap- 
parently a normal  season,  on  April  1 at  Paso  Robles,  California, 
with  a day-degree  accumulation  of  1421  from  the  first  of  January9; 
in  1902  the  blossoming  at  Pomona,  California,  was  on  April  5 with 
an  accumulation  of  over  2300  day-degrees10  and  in  1903  on  April 
15  with  an  accumulation  of  about  2273  day-degrees.11 

Varietal  Differences. — Of  the  varieties  for  which  data  are 
available  for  all  the  years  of  record,  Minnesota,  Fameuse  and  Pri- 
mate are  the  earliest  blossoming;  Rome,  Ralls  and  Ingram  the 
latest.  Data  are  presented  in  Table  8 showing  the  coefficients  of 
variability  in  summations  to  blossoming  in  these  varieties  from  dif- 


Table  8. — Variability  in  Day-Degree  Summations  From  Various  Dates  to 
Blossoming  in  the  Apple  at  Columbia,  Mo.,  Computed  From 
Maximum  Temperatures  Above  43°F. 


Oct.  1 

Nov.  1 

Dec.  1 

Jan.  1 

Feb.  1 

Feb.  15 

Mar.  1 

Mar.  15 

Apr.  1 

Early  blos- 

soming 

Minnesota 

7.82 

8.95 

8.96 

9.41 

8.98 

8.21 

8.68 

16.78 

Fameuse 

7.91 

8.21 

8.32 

9.15 

9.15 

7.99 

7.69 

13.45 

Primate 

7.48 

8.81 

9.28 

10.90 

10.24 

9.51 

10.41 

15.56 

Av. 

7.77 

8.66 

8.85 

9.75 

9.46 

8.57 

8.93 

15.26 

Weather 

Late  blos- 

8.33 

11.87 

16.58 

18.11 

20.89 

20.09 

22.46 

18.41 

soming 

Rome 

7.61 

9.27 

9.27 

10.32 

10.49 

10.18 

9.82 

8.41 

29.65 

Ralls 

7.84 

9.01 

10.42 

10.14 

11.21 

11.12 

11.23 

8.69 

23.79 

Ingram 

7.58 

9.01 

7.68 

8.15 

8.63 

9.60 

8.76 

9.62 

27.74 

Av. 

7.68 

9.10 

9.12 

9.54 

10.11 

10.29 

9.94 

8.91 

27.06 

Weather 

6.65 

8.50 

9.46 

9.37 

10.62 

9.89 

9.88 

7.01 

15.15 

Relation  of  Temperature  to  Blossoming — Apple  and  Peach  21 


ferent  dates  on  the  43°  maximum  basis.  The  variability  in  the 
weather  to  average  data  of  blossoming  as  compared  with  the 
Ohio  figures  is  generally  greater  in  the  early  blossoming  varie- 
ties and  lower  in  the  late  blossoming.  Much  of  this  difference  may 
be  attributed  to  the  smaller  number  of  years  considered,  since  1912 
was  marked  by  great  deficiency  in  temperature  until  near  the  av- 
erage date  of  blossoming  for  the  early  varieties,  but  was  more 
nearly  normal  by  the  average  date  for  the  late  blossoming  var- 
ieties. Omission  of  this  year  from  the  record  would  reduce  the 
variability  in  the  summations  to  the  average  date  of  the  early 
blossoming  varieties  very  materially.  The  lower  variability  in 
the  Columbia  figures  to  the  average  date  for  the  late  blossoming 
varieties  may  be  due  to  the  greater  number  of  day-degrees  involved 
or  it  may  be  accidental.  The  probable  error  of  the  mean  from 
January  1,  is,  for  Columbia  —40,  as  compared  with  —21  for  Wau- 
seon. 

As  they  stand,  the  figures  in  Table  8 show,  though  not  at 
all  clearly,  the  same  general  tendencies  in  the  early  blossoming 
varieties  as  those  appearing  in  the  Wauseon  data,  with  the  ap- 
parently significant  date  earlier.  Those  for  the  late  blossoming 
varieties,  however,  show  no  agreement  greater  than  that  in  the 
weather  to  their  average  date  of  blossoming.  The  drop  to  8.91 
on  March  15  might  be  significant  were  it  not  for  the  even  lower 
figure  (7.01)  for  the  weather  check.  Though  the  low  value  of  the 
latter  is  obviously  accidental,  it  precludes  the  attachment  of  any 
significance  to  the  former. 

Another  way  of  comparing  these  two  groups  of  apple  varieties 
is  through  coefficients  of  correlation  between  accumulations  and 
the  date  of  blossoming,  somewhat  after  the  manner  used  by  Aoki 
and  Tazika4  in  the  sweet  cherry.  In  this  case  any  relationship 
would  be  shown  by  a negative  correlation.  As  shown  in  Table  9 
the  correlation,  wherever  there  is  one,  is  stronger  in  the  early 

Table  9. — Coefficients  of  Correlation  Between  Heat  Accumulations  (Above 
43°,  Max.)  and  Date  of  First  Blossoms  in  Apple  at  Columbia,  Mo. 


Period  of  Early  blossoming  Late  blossoming 

accumulation  varieties  varieties 


January  0.174±0.13  0.168±0.18 

February  — 0.369±0.16  — 0.142±0.11 

March  — 0.856±0.05  — 0.510±0.14 

February  15— March  15  — 0.473±0.15  —0.065 


22 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


blossoming  varieties.  Since  the  time  interval  between  the  per- 
iods of  accumulation  considered  and  the  blossoming  is  shorter  in 
the  early  blossoming  than  in  late  blossoming  varieties,  there  is 
less  opportunity  for  disturbing  variations  in  the  unmeasured  in- 
terval and  the  correlation  would  be  expected  to  be  greater  in  the 
former.  However,  even  with  this  allowance,  there  seems  some 
indication  that  the  date  of  effective  temperatures  is  earlier  in  the 
early  blossoming  than  in  the  late  blossoming  varieties. 

Different  Temperature  Basis. — Since  it  seems  possible  that 
the  late  blossoming  of  some  varieties  may  be  due  to  lack  of  re- 
sponse to  certain  temperatures  which  are  effective  with  the  early 
blossoming  varieties,  variability  coefficients  based  on  a higher 
minimum,  50°,  are  presented  in  Table  10.  Here,  curiously  enough 
in  view  of  the  Wauseon  results,  the  variability  for  the  early  blos- 
soming varieties  is  generally  decreased,  though  the  variability  of 
the  weather  is  increased.  The  full  significance  of  this  is  not  clear 
though  the  study  of  the  records  for  single  years  which  follows 
may  explain  it  in  part.  In  the  late  blossoming  varieties  the  varia- 
bility in  summations  to  blossoming  generally  decreases  somewhat, 
while  that  of  the  summations  to  the  average  date  increases.  The 
changes  are  too  slight,  however,  to  be  indicative.  One  possibly 
significant  change  is  in  the  figures  for  March  15  where  the  vari- 
ability increases  enough  to  give  the  tangent  a value  of  0.7034.  Of 
itself  this  is  not  sufficiently  low  to  have  much  weight,  but  in  con- 


Table  10. — Variability  in  Day-Degree  Summations  From  Various  Dates  to 
Blossoming  in  the  Apple  at  Columbia,  Mo.,  Computed  From  Maximum 
Temperatures  Above  50° F. 


Nov.  1 

Dec.  1 

Jan.  1 

Feb.  1 

Feb.  15 

Mar.  1 

Mar.  15 

Apr.  1 

Early  blossoming 

Minnesota 

10.55 

8.45 

8.31 

8.59 

9.81 

10.59 

15.96 

Fameuse 

10.04 

8.78 

9.47 

8.61 

7.45 

6.92 

13.37 

Primate 

9.42 

9.38 

8.95 

7.81 

8.41 

8.30 

13.12 

Av. 

10.00 

8.87 

8.91 

8.34 

8.56 

8.60 

14.15 

Weather 
Late  blossoming 

14.44 

20.69 

21.45 

24.22 

24.09 

25.86 

23.17 

Rome 

11.51 

9.34 

8.96 

9.50 

9.34 

8.94 

8.42 

34.14 

Ralls 

10.10 

10.59 

9.94 

10.65 

9.93 

9.33 

8.10 

27.76 

Ingram 

10.61 

8.69 

7.72 

8.46 

9.36 

8.89 

10.65 

31.73 

Av. 

10.74 

9.54 

8.87 

9.54 

9.54 

9.05 

9.06 

31.21 

Weather 

8.73 

9.80 

8.28 

10.10 

10.81 

10.66 

12.88 

21.07 

Relation  of  Temperature  to  Blossoming — Apple  and  Peach  23 


nection  with  the  condition  shown  for  this  date  in  Table  8 it  may 
have  some  meaning. 

Seasonal  Differences. — Some  interesting  weather  variations 
with  related  responses  are  shown  by  the  graphs  of  yearly  accum- 
ulations shown  in  figures  2,  3 and  4.  These  are  grouped  more 
or  less  at  random,  the  chief  aim  being  to  present  the  years  of 
peach  blossoming  in  two  diagrams. 

The  first  four  years  of  the  record  are  shown  in  figure  2.  Two 
of  the  four,  1906  and  1907,  were  rather  high  in  accumulations  to 
March  15  and  diverged  widely  from  that  time ; the  1905  curve 


Fig.  2. — Accumulations  (in  day-degrees)  to  blossoming  at  Columbia,  Mo. 


shows  a markedly  low  winter  accumulation  followed  by  rapid  ad- 
vance; 1908  is  noteworthy  for  steadiness  of  the  accumulation  from 
March  1.  Another  interesting  relationship  is  the  identity  of  ac- 
cumulations about  March  15  of  the  two  pairs  of  curves.  The 
agreement  in  these  pairs  in  the  accumulations  to  blossoming  for 
Rome  and  the  agreement  in  summations  from  March  15  are  re- 
markably close.  The  tangent  in  this  case  is  0.244.  The  agreement 
in  Primate  is  even  closer,  the  tangent  being  in  fact  0.222,  but  the 
agreement  is  not  within  the  same  pairs  as  in  Rome.  In  both  cases 


24 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


of  blossoming  at  the  lower  summation  the  accelerating  influence 
of  high  temperature  is  apparent  in  the  steepness  of  the  gradient. 

The  1906  and  1908  curves  are  rather  close  to  parallel  for  some 
time  and  blossoming  of  Primate  occurs  at  the  same  level  on  them. 
The  Elberta  peach  blossoms  at  a lower  level  on  the  1908  curve. 
These  curves  crossed  about  March  10  and  accumulations  then 
were  identical,  but  the  more  rapid  rise  from  that  point  in  1908 
evidently  had  more  effect  on  Elberta  than  on  Primate.  Compari- 
son of  the  1905  and  1908  curves  indicates  the  effect  of  the  sharp 
rise  after  March  15  in  hastening  the  development  of  the  Primate 
blossoms. 

In  1905  the  Primate  apple  blossomed  ahead  of  the  Elberta 
peach.  The  arrangement  in  the  figure  shows  that  this  was  due  to 
Elberta  being  late  in  blossoming  rather  than  Primate  being  early. 
This  was  the  year  of  very  little  accumulation  until  after  March  1. 
Apparently  the  cold  weather  held  the  peach  dormant  until  high 
temperatures  could  become  effective  on  the  apple,  as  in  Ohio  in 
1912,  shown  in  figure  1.  It  should  be  stated,  however,  that  a 
considerable  amount  of  winter-killing  of  buds  occurred  during  the 
winter  of  1904-1905  and  that  the  blossoming  of  Elberta  as  re- 
corded is  doubtless  later  than  it  would  have  been  with  a full  crop, 
since  generally  the  more  advanced  buds  are  more  readily  killed. 
Furthermore,  Chandler13  mentions  a mild  form  of  winter  injury 
which  retards,  but  does  not  prevent  blossoming.  Even  with  this 
allowance,  however,  the  closeness  of  Elberta  and  Primate  is  in- 
dicative of  the  influences  mentioned. 

It  is  interesting  that  Morgan26  working  at  Ithaca,  New  York, 
reported  the  apple  to  start  development  earlier  in  the  spring  than 
the  peach  but  that  the  peach  rapidly  overtook  it.  This  might 
well  be  the  case  if  the  investigation  were  carried  on  in  such  years 
as  1905  or  where  the  common  season  resembles  the  1905  season. 
On  the  other  hand  Drinkard14  in  Virginia  reported  more  ad- 
vancement during  the  winter  in  the  peach  than  in  the  apple.  As- 
suming the  peach  to  require  higher  temperatures  than  the  apple 
it  might  start  later  than  the  apple  in  seasons  that  are  cool,  but 
with  warm  temperatures  it  starts  before  the  apple. 

In  figure  3 are  presented  curves  for  the  remaining  years  for 
which  peach  blossoming  dates  are  available,  with  that  for  1912 
added  for  comparison.  The  successive  spring  freezes  of  1921  de- 
stroyed apple  blossoms  so  extensively  that  blossoming  records 
were  not  taken,  consequently  the  curve  for  that  year  is  not  car- 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  25 


ried  beyond  the  blossoming  of  Elberta,  which  occurred  before  any 
damage  had  been  inflicted  and  is  therefore  reliable.  At  first  glance 
the  high  accumulations  for  Elberta  in  1909  and  1911  are  outstand- 
ing and  apparently  inconsistent.  These  years,  however,  were  char- 
acterized by  a considerable  amount  of  winter  killing  of  buds,  the 
damage  in  1909  in  Elberta  at  Columbia  amounting,  according  to 
Chandler,12  to  97.3  per  cent.  Data  are  not  available  on  the  ex- 
tent of  the  damage  to  Elberta  in  1911,  but  since  it  ranged  from 


T* 

4 


os 


< 


t 


ci 

G, 


*< 


r~< 

I 


Fig.  3. — Accumulations  (in  day-degrees)  to  blossoming  at  Columbia,  Mo. 


29.8  per  cent  to  79  per  cent  in  other  varieties,  it  must  have  been 
considerable.  These  dates,  then,  represent  the  opening  of  only 
a very  small  portion  of  the  total  number  of  blossoms  and  these, 
presumably,  those  that  were  least  advanced  during  the  winter  and 
would  be  the  last  to  open  in  the  spring.  With  these  allowances, 
the  line  connecting  the  blossoming  dates  of  Elberta  would  become 
nearly  horizontal,  signifying  a rather  close  agreement  in  totals  to 
blossoming. 

In  Rome  the  differences  in  day-degrees  at  blossoming  are  in 
the  same  order  as  the  differences  on  March  15  with  one  exception. 


26 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


This  is  on  the  1909  curve  where  Rome  seems  unduly  late  in  blos- 
soming. Since  yield  records  are  not  available  for  this  variety  the 
amount  of  bloom  this  year  cannot  be  stated.  That  Rome  was  “out 
of  step”  in  this  case  is  shown  by  the  fact  that  this  was  the  only 
year  in  the  record  when  Ingram  blossomed  ahead  of  Rome.  If 
this  were  due  to  scarcity  of  crop  so  that  the  only  blossoms  ap- 
pearing were  terminals — as  is  sometimes  the  case — this  discrep- 
ancy would  be  explained.  However,  even  with  this  allowance, 
the  agreement  is  very  little  greater  in  summations  from  March 
15  to  May  1. 


5 to  0£  ei  >• 

4 £ £ 1 i 

Fig.  4. — Accumulations  (in  day-degrees)  to  blossoming  at  Columbia,  Mo. 

In  figure  4 are  shown  graphs  for  years  in  which  no  blossom- 
ing dates  for  the  Elberta  peach  are  available;  to  these  1907  is 
added  because  of  its  close  similarity  to  1918  after  March  1.  These 
graphs  are  strikingly  similar,  excepting  the  1912  curve.  In  1913, 
1916  and  1917  Primate  appears  to  have  been  retarded  by  a week 
of  cool  weather  in  early  April,  though  its  extreme  lateness  in  1917 
can  be  only  partly  explained  in  this  way.  Here  again,  sparseness 
of  the  crop  may  be  a partial  explanation,  since  study  of  the  spurs 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  27 


of  this  tree  in  the  orchard  shows  very  little  blossoming  in  that 
year.  The  blossoming  of  Rome  at  a comparatively  low  accumula- 
tion in  1913  may  be  explained  by  the  rapid  rise  in  temperature 
subsequent  to  April  15. 

The  evident  retarding  effect  of  the  cold  weeks  in  early  April 
in  1913,  1916  and  1917  and  the  absence  of  influence  of  these  weeks 
on  Rome,  though  it  is  clearly  responsive  to  high  temperature,  raises 
an  interesting  question.  Since  the  buds  of  Primate,  which  was 
retarded,  were  more  advanced  at  these  periods  than  those  of  Rome, 
which  apparently  was  not  retarded,  it  seems  quite  possible  that 
optimum  temperatures  vary  as  the  buds  advance  toward  opening. 
The  decrease  in  the  rate  of  the  temperature  rise  in  1907  after 
Primate  was  in  blossom  and  Rome  presumably  well  advanced, 
seems  to  have  had  a retarding  effect. 

There  is  a rather  strong  trend  toward  uniformity  in  summa- 
tions to  blossoming  in  these  curves,  if  1912  be  disregarded.  This 
does  not,  however,  necessarily  signify  that  temperatures  of  Jan- 
uary and  February  are  effective,  since  the  accumulations  are  very 
much  alike  on  March  1 or  even  on  March  15.  Here  again,  plot- 
ting the  curves  with  March  1 as  the  starting  point  or,  for  Rome, 
March  15,  secures  much  greater  agreement. 

Considering  all  graphs  shown,  and  making  the  allowances  in- 
dicated, there  is  a rather  marked  tendency  for  uniformity  in  sum- 
mations from  January  1 to  blossoming,  in  the  Elberta  peach. 
Where  the  uniformity  appears  in  the  blossoming  of  the  apples  it 
is  accompanied  by  an  approximate  uniformity  in  the  accumula- 
tions at  some  date  subsequent  to  January  1.  Nowhere,  however, 
is  there  clear  evidence  pointing  to  uniformity  or  difference  in  the 
end  of  the  rest  period  between  the  early  and  the  late  blossoming 
apples.  The  graphs  in  figure  3,  contrasting  warm  springs  with 
a very  cold  spring,  suggest  a difference  in  the  rest  period. 

In  Table  11  are  assembled  data  showing  the  fluctuating  dif- 
ference between  30  varieties  of  apple  for  all  the  years  of  record. 
Included  in  this  are  all  the  varieties  for  which  data  are  complete; 
in  most  cases  the  records  are  from  the  same  trees  throughout. 
The  seasonal  difference  in  dates  of  first  bloom  (range)  is  shown 
to  vary  from  5 to  27  days  and  the  average  deviation  from  0.97  to 
4.75  days.  That  this  difference  between  the  first  and  last  blos- 
soming variety  is  little  more — or  is  even  less — constant  when  ex- 
pressed in  terms  of  heat  is  shown  by  comparison  of  the  maximum 
range  in  day-degrees  (519)  with  the  minimum  (187).  The  gen- 


28 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


Table  11. — Variations  in  Blossoming  Among  Thirty  Apple  Varieties  at 

Columbia,  Mo. 


Days 

Day-degrees 

Aver. 

date 

Duration 
of  bloom 

Aver. 

devi- 

Aver. 

accumu- 

Aver. 

devi- 

Aver. 

daily 

Year 

of 

bios. 

Range 

in  earliest 
variety 

ation 

lation 

Range 

ation 

acc.  dur- 
ing bios. 

1905 

99.6 

24 

22 

2.85 

898.7 

519 

69.5 

21.6 

1906 

111.2 

13 

12 

1.69 

926.4 

415 

51.1 

31.9 

1907 

87.9 

22 

21 

3.14 

886.8 

323 

48.4 

14.7 

1908 

101.4 

14 

12 

1.87 

943.6 

342 

45.1 

24.4 

1909 

111.2 

18 

13 

3.32 

1191.3 

419 

52.3 

23.3 

1911 

104.7 

18 

20 

2.89 

1109.8 

380 

61.4 

21.1 

1912 

115.7 

8 

16 

1.25 

766.0 

193 

30.0 

24.1 

1913 

109.5 

12 

12 

1.80 

964.9 

315 

48.6 

26.3 

1914 

111.2 

11 

10 

1.59 

976.1 

359 

51.7 

32.6 

1915 

111.0 

5 

6 

0.97 

879.6 

187 

36.4 

37.4 

1916 

107.9 

13 

13 

1.47 

1059.1 

279 

40.3 

21.5 

1917 

111.7 

20 

8 

2.81 

1100.0 

313 

63.4 

15.6 

1918 

100.3 

27 

33 

4.75 

1087.3 

418 

69.1 

15.5 

Av. 

106.4 

15.8 

14.5 

2.34 

983.8 

343 

51.3 

23.8 

eral  accelerating  effect  of  high  temperature  is  evident  in  the  av- 
erage of  the  daily  temperature  accumulations  during  the  three 
shortest  ranges,  31.3,°  and  during  the  three  longest,  17.3.° 

No  constant  on  the  basis  used  here  will  measure  the  dif- 
ference between  blossoming  in  the  earliest  apple  and  the  latest. 
It  is  quite  likely  that  an  exponential  or  a “physiological”  system 
would  measure  this  brief  span  more  closely.  It  is,  however,  quite 
as  probable  that  conditions  before  the  blossoming  of  the  earliest 
apples  vary  from  season  to  season  and  that  this  event  may  find 
the  late  blossoming  variety  at  various  stages.  When  the  early 
blossoming  variety  is  not  held  back  by  unfavorable  weather  the 
late  blossoming  kind  will  lag  behind ; when  the  early  blossoming 
variety  is  retarded  the  difference  will  be  less,  other  things  equal. 
This  indicates  a difference  in  date  of  effective  temperatures.  How- 
ever, it  is  noteworthy  that  no  matter  how  much  the  season  is  re- 
tarded and  how  small  the  range  between  the  varieties,  the  early- 
blossoming  kinds  bloom  first  and  the  late-blossoming  varieties 
bloom  last.  This  indicates  a difference  in  temperature  require- 
ments. 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  29 


Leaf  and  Fruit  Buds  Compared. — Apparently  the  opening  of 
blossoms  and  the  unfolding  of  leaves  respond  somewhat  differ- 
ently to  a given  set  of  conditions.  Bailey5  says  that  in  the  southern 
states  plum  flowers  “tend  to  appear  wholly  in  advance  of  the 
leaves,  and  they  are  borne  upon  short  stalks,  or  may  be  nearly 
or  quite  sessile.  In  the  North,  the  flowers  and  leaves  are  gen- 
erally coetaneous,  and  the  flower  stalks  are  usually  longer.”  Bal- 
lard and  Volck6  report  that  spraying  with  nitrate  of  soda  in  Feb- 
ruary hastened  the  opening  of  flowers  but  not  of  leaf  buds,  in 
apples  and  pears.  Table  12  shows  the  variability  in  summations 
to  appearance  of  the  first  fully  formed  leaf  at  Wauseon,  Ohio. 
Since  the  period  of  record  is  not  identical  with  that  for  blossom- 
ing the  figures  are  not  strictly  comparable.  However,  the  dif- 
ferences between  the  values  of  the  tangents  in  Tables  4 and  12 
seem  considerable  enough  to  signify  some  difference  between 
leaves  and  blossoms  in  their  responses.  In  some  years  blossoms 
preceded  leaves;  other  years  showed  the  opposite  condition. 

Table  12. — Variability  in  Day-Degree  Summations  (Maximum,  Above  43°) 
to  Date  oe  the  Appearance  oe  the  First  Fully  Formed  Leae  in  the 


Apple  at  Wauseon,  Ohio. 

Variability  in 

Summations  Variability  in  summations  to  Tangent 

beginning  summations  average  date 


September  1 7.30  7.47  0.98 

October  1 9.05  10.23  0.88 

November  1 10.21  15.74  0.65 

December  1 11.74  17.69  0.66 

January  1 11.90  18.60  0.64 

February  1 11.28  17.73  0.64 

March  1 11.41  17.63  0.65 

April  1 16.42  19.09  0.86 


The  data  for  Columbia  record  a somewhat  different  phase  of 
vegetative  development,  namely,  the  opening  of  the  leaf  buds. 
Very  rare  indeed  in  these  records  is  the  case  where  the  opening 
of  the  first  blossom  precedes  the  opening  of  the  first  leaf  bud ; al- 
most invariably  the  leaf  buds  open  before  the  blossoms.  The  mar- 
gin of  difference  varies,  however.  In  three  typical  early  blossom- 
ing varieties  the  average  difference  for  13  years  is  7 days;  in 
three  typical  late  blossoming  varieties  it  is  11  days.  The  late 
blossoming  varieties  blossomed  on  the  average  13  days  after  the 


30 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


opening  of  the  first  bloom ; their  leaves  appeared,  on  the  average, 
10  days  after  the  opening  of  the  first  leaf  bud. 

Evidence  from  Microscopic  Examination. — Explanation  of 
much  of  the  lack  of  agreement  among  the  variability  coefficients  of 
the  several  varieties  represented  in  Tables  8 and  10  is  found 
through  microscopic  examination  of  flower  buds  at  various  times 
during  the  winter.  Typical  photomicrographs  of  preparations  made 
by  Mr.  V.  R.  Boswell  are  shown  in  Plates  I,  II  and  III.  Oldenburg 
and  Primate  represent  the  earliest  blossoming  varieties;  Rome, 
Daru  and  Cilligos  the  latest.  The  two  last  are  included  since 
they  have  been  used  extensively  in  the  apple  breeding  work  of 
the  Missouri  Station.  Daru  blossoms  at  about  the  same  time  as 
Ingram ; Cilligos  is  the  latest  blossoming  of  all  varieties  under 
observation. 

Plate  I shows  the  stages  reached  by  several  varieties  on  Feb- 
ruary 2,  1920.  Oldenburg  is  clearly  more  advanced  than  the  other 
varieties.  In  the  other  cases,  the  correspondence  between  develop- 
ment on  this  date  and  the  order  of  blossoming  is  not  so  close. 
Fameuse,  the  second  earliest  in  blossoming,  is  no  farther  advanced 
than  Daru,  the  second  latest  in  blossoming.  Gano  and  York,  mid- 
season varieties,  are  apparently  at  the  same  stage  as  Cilligos,  the 
latest  of  all. 

In  Plate  II  are  shown  the  stages  on  three  dates,  November 
2,  1921,  January  28,  1922,  and  February  20,  1922,  for  three  varie- 
ties, Oldenburg,  Primate  and  Wealthy.  The  first  two  are  dis- 
tinctly early  in  blossominng;  Wealthy  might  be  classed  either 
among  the  last  of  the  early  blossoming  or  among  the  earliest  of 
the  mid-season  varieties.  Here  the  advancement  of  Oldenburg  in 
November  is  marked;  this  appears  clearly  to  be  a factor  in  its 
early  blossoming  since  subsequent  samples  show  relatively  slight 
development.  The  other  early  blossoming  variety,  Primate,  shows 
a quite  different  condition.  Its  November  stage  is  not  advanced; 
indeed,  Daru,  one  of  the  latest  blossoming,  shows  equal  or  greater 
pistil  development  on  this  date.  Its  changes  through  the  winter, 
however,  are  notable  and  suggest  that  this  variety  has  a factor 
producing  early  blossoming  quite  different  from  or  more  intense 
than  that  evident  in  Oldenburg.  Wealthy,  equally  or  more  ad- 
vanced in  early  November,  does  not  develop  as  rapidly  through 
the  winter. 

Plate  III  records  the  development  of  the  buds  in  three  late 
blossoming  varieties  sampled  on  the  same  dates  as  those  of  the 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  31 


early  blossoming  varieties  shown  in  Plate  II.  Daru,  the  second 
latest  in  blossoming  of  all  the  varieties  shown,  is  among  the  more 
advanced  on  November  2.  Its  lateness  is  due  apparently  to  its 
lack  of  responsiveness  to  temperatures  with  which  Primate  de- 
velops. Rome  presents  an  anomaly  in  that  it  is  perhaps  the  least 
advanced  in  November  and  apparently  advances  little  or  none  to 
February  20;  nevertheless  it  comes  into  blossom  ahead  of  Daru 
and  Cilligos. 

The  winter  of  1921-1922  in  Columbia  was  mild  in  the  sense 
that  there  was  little  very  cold  weather.  However,  as  measured  in 
day-degrees  above  43°  it  was  not  warmer  than  the  ordinary  sea- 
son; the  monthly  accumulations  from  November  to  February  in- 
clusive being  respectively  345,  167,  92  and  178.  November  accum- 
ulations were  below  the  average  (426)  and  December  above  (39), 
January  somewhat  below  the  average  (122)  and  February  some- 
what above  (139).  The  samples  shown  here,  however,  were  gath- 
ered on  February  20,  when  the  accumulation  for  the  month  had 
reached  102  day-degrees  only  and  before  the  warmest  weather 
of  the  month.  Consequently  such  development  as  is  shown  to  be 
connected  with  temperature  for  this  winter  may  be  regarded  as 
normal  for  this  locality. 

Evidently,  then,  early  blossoming  in  apples  involves  at  least 
two  factors : first,  the  stage  of  advancement  reached  at  the  ap- 

proach of  winter,  as  exemplified  by  Oldenburg;  second,  ability  to 
develop  through  the  winter,  as  shown  by  Primate.  Late  blossom- 
ing, presumably,  is  due  to  the  absence  of  both  these  factors  or  to 
the  presence  of  strong  inhibitors  of  the  second.  The  ideal  late 
blossoming  variety  as  represented  by  Cilligos  is  backward  in  de- 
velopment in  the  fall  and  advances  little  through  the  winter.  It 
is  plausible  that  mixed  inheritance  of  these  factors  gives  the  mid- 
season blossoming  shown  by  the  majority  of  commercial  var- 
ieties, though  Daru  appears  to  have  one  factor  for  earliness  despite 
its  late  blossoming.  This  seems  the  more  likely  since  its  crosses 
with  Delicious  now  growing  in  the  Experiment  Station  grounds 
include  only  very  few  late  blossoming  varieties,  a smaller  per- 
centage than  those  shown  by  the  majority  of  the  crosses  involving 
Ingram,  another  late  blossoming  variety. 

The  behavior  of  the  late  blossoming  varieties  indicates  either 
a requirement  of  higher  temperatures  for  advancement  or  the 
temporary  presence  of  a development-inhibiting  factor  that  is  ab- 
sent in  the  early  blossoming  kinds.  If  late  blossoming  is  due  to 


32 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


a higher  temperature  requirement,  the  difference  between  late 
and  early  blossoming  kinds  should  be  diminished  by  forcing  in  a. 
greenhouse.  If  late  blossoming  is  due  to  the  persistence  of  the 
rest  period  in  some  form  these  differences  should  decrease  as  the 
season  advances.  Table  13  shows  the  results  obtained  by  forc- 
ing twigs  of  Primate  (hypothetically  without  or  over  the  rest 
period  )and  of  Rome  (hypothetically  still  in  the  rest  period).  The 
stages  observed  in  the  two  varieties  differ,  but  comparison  is  pos- 
sible. Though  the  buds  started  March  3 were  kept  in  a cooler 
house  than  that  used  for  the  two  earlier  lots,  enough  cooler  ap- 
parently to  retard  Primate,  Rome  started  in  a shorter  time.  The 
lower  temperatures  actually  retarded  the  early  blossoming  variety 
more  than  the  late  blossoming.  This,  with  the  progressive  short- 
ening of  the  period  of  forcing  in  Rome,  indicates  the  rest  period 
as  a factor  rather  than  a differential  temperature  requirement. 


Table  13. — Number  oe  Days  Involved  in  Forcing  Blossom  Buds  oe  Primate 
and  Rome  Apples,  1922. 


Date  forcing  started  Days  to  blossoms  open  Days  to  buds  starting 

in  Primate  in  Rome 

February  16  17  15 

February  25  14  14 

March  3 20  13 


Comparison  of  Plates  IV  and  V shows  that  the  difference  be- 
tween varieties  are  greater  when  they  are  forced  in  the  green- 
house than  when  the  buds  develop  in  the  orchard.  This  points 
in  the  same  direction  as  the  evidence  just  cited. 

Other  Considerations. — Analysis  of  the  records  of  42  trees 
for  which  data  are  complete  shows  no  relation  between  the  date 
of  terminal  bud  formation  on  shoots  and  the  date  of  spur  blossom- 
ing in  the  spring,  the  correlation  coefficient  being  0.085—0.04.  It 
is  possible,  however,  that  comparison  of  trees  under  different 
cultural  conditions  might  show  a relation  of  this  sort,  though  it 
is  doubtful  if  it  should  not  be  considered  an  associated  rather  than 
a causative  condition. 

Incidentally  the  relation  to  cross  pollination  of  differences  in 
blossoming  may  be  mentioned.  Comparison  of  the  figures  in  the 
column  headed  “Range”  with  those  in  that  headed  “Duration  of 
bloom  in  earliest  variety”  in  Table  11  shows  that  in  eight  years 
of  the  thirteen  recorded  the  earliest  variety  was  out  of  bloom  be- 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  33 


fore  the  latest  blossoming  came  in.  In  two  years  the  date  of  last 
blossom  in  the  one  and  of  first  bloom  in  the  other  were  identical. 
In  one  only  was  the  overlapping  sufficient  to  ensure  abundant 
cross  pollination.  In  this  section,  then,  when  very  early  blossom- 
ing kinds  are  planted  with  very  late  blossoming  kinds,  cross  pol- 
lination can  be  ensured  only  by  a third  variety,  intermediate  in 
blossoming  season.  This  will  provide  pollination  for  the  early 
blooming  kinds  with  its  first  blossoms  and  for  the  late  blooming 
with  its  last  blossoms.  Most  of  the  commercial  varieties  grown 
in  Missouri  fall  into  the  intermediate  class  in  blossoming  and 
may  be  counted  on  with  safety  so  far  as  cross  pollination  is  con- 
cerned. However,  it  is  possible  that  the  reputation  of  the  Rome 
for  light  bearing  in  Missouri,  though  in  Ohio  it  has  not  met  that 
objection,  is  due  to  the  greater  extent  of  the  blossoming  season 
in  Missouri  so  that  Rome  may  in  some  seasons  be  in  bloom  alone 
while  in  Ohio  the  difference  ordinarily  would  be  less  marked. 

Table  14  shows  the  blossoming  dates  of  several  peach  va- 
rieties, selected  to  permit  comparison  with  dates  for  the  same 
varieties  at  points  with  winters  considerably  milder  than  those  at 
Columbia.  For  compactness  these  are  expressed  in  days  of  the 
year  rather  than  of  the  month.  Though  the  list  for  most  years 
at  Columbia  is  more  extensive  than  those  given  for  Alabama  or 
California,  the  range  in  blossoming  represented  is  less  in  every 
case.  In  other  words,  just  as  the  blossoming  of  the  apple  in  dis- 
tinctly cold  sections  has  a narrower  range  than  at  Columbia,  so 
the  peach  at  Columbia  has  a narrower  range  than  at  points  farther 
south.  Cool  weather  during  the  peach  blossoming  season  at  Au- 
burn, Alabama,  may  have  prolonged  the  season  of  1911  to  an  un- 
usual length,  but  the  normal  blossoming  range  of  the  varieties 
named  in  this  paper  is  apparently  as  great  or  greater  than  the  maxi- 
mum recorded  for  Columbia.36  The  range  shown  for  Pomona, 
California,  is  apparently  normal  for  that  point. 

The  slight  difference  between  all  varieties  at  Columbia  in 
1907,  the  year  of  earliest  blossoming  for  which  an  approximately 
complete  record  is  available,  indicates  that  the  rest  period  as  a 
factor  in  the  date  of  blossoming  in  the  peach  is  not  operative  here. 
This  was  a year  of  rather  high  temperature  from  January  on.  A 
considerably  greater  number  of  varieties  than  is  here  reported 
showed  almost  as  close  agreement  in  blossoming  in  1921,  when 
the  season  was  even  earlier  than  that  of  1907.  Any  differences  in 
the  rest  period  which  might  be  concealed  by  the  retarding  effect 


34 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


of  an  ordinary  winter  on  the  earliest  varieties  should  become  evi- 
dent in  these  seasons  of  exceptionally  high  late  winter  tempera- 
tures, as  soon  as  growth  is  possible.  The  third  year  of  closeness 
in  blossoming,  1906,  was  characterized  by  rather  low  accumulation 
during  winter,  with  a rapid  advance  about  the  time  of  blossom- 
ing. The  spread  of  the  year  of  greatest  range  is  due  apparently 
to  unequal  winter-killing  of  blossoms  and  to  the  slow  accumu- 
lation of  temperature,  which  brought  out  minor  differences  in  re- 
sponse to  heat  or  merely  delayed  the  opening  of  those  varieties 
which  had  fewest  buds.  In  warmer  climates  it  seems  quite  pos- 
sible that  these  differences  in  blossoming  are  due  to  differences  in 
the  termination  of  the  rest  period,  particularly  since  the  Peen-to 
peaches  there  blossom  much  earlier  than  those  recorded  in  Table 
14,  and  almonds  in  January  or  February. 


Table  14— Peach  Blossoming  Dates  (In  Days  oe  Year)  at  Various  Points. 


Variety 

Columbia, 

Mo. 

Au- 

burn 

Ala. 

Pomona 

Calif. 

1905 

1906  1907 

1908 

1909  1911 

1914 

1915 

1921 

1911 

1894 

1895 

1902  1903 

Alexander 

98 

102 

82 

90 

101 

95 

78 

76 



8S 

Briggs  Red 

98 

103 

83 

90 

96 

92 

78 

63 



91 

Carman 

98 

102 

83 

85 

— 

95 

98 

107 

76 

52 

Champion 

97 

103 

82 

86 

94 

94 

99 

107 

76 

52 

Chinese  Cling 

100 

102 

83 

86 

96 

93 

97 

106 

75 

46 

71 

64 

84 

77 

Crawford  Early 



102 

82 

80 

96 

93 

96 

108 

— 

35 

74 

62 

91 

84 

Crawford  Late 

99 

102 

83 

86 



96 

94 

72 

60 

91 

69 

Elberta 

98 

102 

82 

85 

95 

92 

97 

107 

74 

45 

Family  Favorite 

99 

102 

82 

85 

95 

93 

44 

Foster 

101 

102 

82 



96 

95 

100 

109 





72 

63 

95 

74 

Globe 



103 

82 

86 

96 

95 

46 

Heath  Cling 

97 

104 

82 

84 

95 

96 

103 

72 

66 

91 

69 

Henrietta 

102 

103 

82 

88 

97 

65 



— 

Lemon  Cling 



102 

83 

88 

96 

93 

103 

105 

73 

— 



Mayflower 

95 

100 

105 

— 

57 

Mountain  Rose 

98 

107 

75 

— 

76 

60 

91 

77 

Oldmixon  Cling 

98 

102 

82 

86 

96 

97 

94 

108 

— 

— 

72 

63 

94 

87 

Oldmixon  Free 

98 

102 

m 

85 

95 

95 

97 

84 

66 

95 

74 

Salway 

98 

103 

83 

85 

96 

92 

103 

107 

76 

46 

73 

66 

61 

91 

Smock 

91 

103 

82 

86 

95 

101 

104 



— 

52 

79 

63 

— 

77 

Sneed 

99 

102 

83 

87 

95 

93 

52 

Susquehanna 

102 

103 

83 

86 

— 

96 

50 

72 

63 

— 

69 

Thurber 

99 

102 

82 

86 

95 

43 

Yellow  St.  John 

98 

103 

82 

86 

99 

96 

72 

63 

64 

79 

Range 

12 

4 

2 

11 

8 

10 

10 

5 

3 

23 

14 

17 

35 

23 

Relation  of  Temperature  to  Blossoming — Apple  and  Peach  35 


SUMMARY 

1.  The  amount  of  heat,  as  measured  in  day-degrees,  received 
by  peaches  from  January  1 to  the  time  of  blossoming,  varies  with 
the  season  and  even  more  with  the  locality. 

2.  The  agreement  in  temperature  accumulations  to  blossom- 
ing from  year  to  year  at  any  one  place  varies  with  the  length  of 
the  time  for  which  they  are  measured  indicating  that  ordinary 
temperatures  are  not  always  effective  or  that  temperature  is  not 
always  a limiting  factor. 

3.  Variability  in  temperature  accumulations  from  various 
dates  to  blossoming  at  Wauseon,  Ohio,  follows  different  orders 
in  the  King  apple  and  the  Late  Crawford  peach,  indicating  that 
the  latter  is  responsive  to  high  temperatures  when  the  former  is 
not. 

4.  The  average  temperature  accumulation  from  January  1 
to  blossoming  in  the  apple  is  somewhat  greater  at  Columbia,  Mo. 
than  at  Wauseon,  Ohio,  but  much  less  than  at  Pomona,  Calif. 

5.  Varietal  differences  in  blossoming  at  Columbia,  Mo.,  in- 
dicate that  the  early  blossoming  varieties  of  apple  become  re- 
sponsive to  ordinary  temperatures  earlier  than  the  late  blossom- 
ing. There  are,  however,  some  inconsistencies  which  are  not  ex- 
plained by  any  mathematical  analysis  attempted. 

6.  Microscopic  examination  of  blossom  buds  indicates  that 
there  are  at  least  two  factors  governing  the  season  of  blossom- 
ing at  Columbia,  Mo.  Oldenburg  blossoms  early  chiefly  because 
the  buds  are  well  advanced  in  the  fall,  Primate  because  the  buds 
develop  through  the  winter.  Daru  is  well  advanced  in  the  fall  but 
does  not  develop  through  the  winter  and  blossoms  la-te.  Cilligos 
is  backward  in  the  fall  and  does  not  advance  through  the  winter ; 
it  is  the  latest  blossoming  variety  observed.  The  mid-season  va- 
rieties apparently  have  a mixed  genetic  constitution  in  this  respect. 

7.  Observations  on  branches  forced  in  the  greenhouse  in- 
dicate that  late  blossoming  is  connected  with  rest  period  influ- 
ences rather  than  with  differential  temperature  requirements. 

8.  Varietal  differences  in  the  peach  at  Columbia  appear  to 
be  masked,  but  may  become  evident  farther  south,  in  the  same 
manner  as  differences  apparent  in  the  apple  at  Columbia  are 
masked  farther  north. 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  37 


ACKNOWLEDGMENTS 

Acknowledgments  are  due  Professors  V.  R.  Gardner,  H.  D. 
Hooker,  Jr.,  and  W.  J.  Robbins  of  the  University  of  Missouri; 
to  Dr.  E.  J.  Kraus  of  the  University  of  Wisconsin,  for  valuable 
aid  and  suggestions;  to  Mr.  George  Reeder  of  the  United  States 
Weather  Bureau,  for  the  temperature  records  for  Columbia;  to 
Mr.  V.  R.  Boswell  of  the  University  of  Missouri,  for  assistance  in 
computations,  for  the  sections  used  in  the  study  of  bud  develop- 
ment and  for  the  photomicrographs ; and  to  those  whose  interest 
and  care  resulted  in  the  accumulation  of  the  phenological  records 
for  Columbia. 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  39 


LITERATURE  CITED 

1.  Abbe,  C.,  U.  S.  D.  A.  Weather  Bur.  Bui.  36  (1905). 

2.  Adanson,  Cited  by  Abbe. 

3.  Angot,  A.,  Cited  by  Abbe. 

4.  Aoki,  S.  and  Tazika.  Y.,  Journ.  Met.  Soc.  Japan.  Apr.  1921,  Abs.  in  U.  S. 

D.  A.  Monthly  Weather  Rev.  49.  609  (1921). 

5.  Bailey,  L.  H.,  The  Evolution  of  Our  Native  Fruits,  p.  199,  N.  Y.  (1898). 

6.  Ballard,  W.  S.  and  Volck,  W.  H.,  Journ.  Agr.  Res.  1 :437  (1914). 

7.  Bigelow,  J.,  Amer.  Jour.  Sci.  1:76  (1820). 

8.  Bradford,  F.  C.,  Oreg.  Agr.  Exp.  Sta.  Bui.  129  (1915). 

9.  Calif.  Agr.  Exp.  Sta.  Ann.  Rept.  1894-1895.  p.  379. 

10.  Calif.  Agr.  Exp.  Sta.  Ann.  Rept.  1902-1903.  pp.  187-190. 

11.  Calif.  Agr.  Exp.  Sta.  Ann.  Rept.  1903-1904.  pp.  175-188. 

12.  Chandler,  W.  H.,  Mo.  Agr.  Exp.  Sta.  Res.  Bui.  8 (1913). 

13.  Chandler,  W.  H.,  Proc.  Am.  Soc.  Hort.  Sci.  12:118  (1915).  — 

14.  Drinkard,  A.  W.,  Jr.,  Va.  Agr.  Exp.  Sta.  Ann.  Rept.  1909-1910.  pp.  159- 

197. 

15.  Fritsch,  K.,  Cited  by  Abbe. 

16.  Gardner,  V.  R.,  Bradford,  F.  C.  and  Hooker,  H.  D.,  Jr.,  Fundamentals 

of  Fruit  Production,  N.  Y.  (1922). 

17.  Hedrick,  U.  P,  N.  Y.  Agr.  Exp.  Sta.  Bui.  407  (1915). 

18.  Ihne,  E.,  Ueber  Beziehungen  zwischen  Pflanzenphanologie  und  Landwirt- 

schaft  p.  9,  Berlin  (1909). 

19.  Johnston,  E.  S.,  Paper  before  Botanical  Society  of  America.  Toronto, 

Ont.  Dec.  29,  1921. 

20.  Linsser,  C.,  Cited  by  Abbe. 

21.  Livingston,  B.  E.,  Physiol.  Res.  1:8  (1916). 

22.  Livingston,  B.  E.  and  Livingston,  G.  J.,  Bot.  Gaz.  56:5  (1913). 

23.  Magness,  J.  R.,  Oreg.  Agr.  Exp.  Sta.  Bui.  139  (1916). 

24.  Merriam,  C.  H.,  U.  S.  D.  A.  Bur.  Biol.  Survey,  Bui.  10  (1898). 

25.  Mikesell,  T.,  U.  S.  D.  A.  Mo.  Weath.  Rev.  Sup.  2 (1915). 

26.  Morgan,  W.  M.,  Thesis.  Cornell  Univ.  1902 — Cited  by  Wiegand,  K.  M., 

Bot.  Gaz.  41:373  (1906). 

27.  Palladin,  V.  J.,  Plant  Physiology.  Eng.  ed.  by  Livingston.  Phila.  (1918). 

28.  Price,  H.  L.,  Va.  Agr.  Exp.  Sta.  Ann.  Rept.  1909-1910.  p.  206. 

29.  Reaumur,  R.  A.  F.  de.,  Mem.  Acad,  des  Sciences.  1735  p.  545.  cited  by 

Abbe. 

30.  Sandsten,  E.  P.,  Wis.  Agr.  Exp.  Sta.  Bui.  137  (1906). 

31.  Schimper,  A.  F.  W.,  Plant  Geography  upon  a Physiological  Basis  p.  37. 

Oxford,  1903. 

32.  Seeley,  D.  A.,  U.  S.  D.  A.  Mo.  Weather  Rev.  45:354  (1917). 

33.  Swingle,  W.  T.,  U.  S.  D.  A.  B.  P.  I.  Bui.  53  (1904). 

34.  Waugh,  F.  A.,  Vt.  Agr.  Exp.  Sta.  Ann.  Rept.  11:270  (1898). 

35.  Weber,  F.,  Sitzungsber.  d.  Akad.  d.  Wiss.  Wien.  125,  330  (1916). 

36.  Williams,  P.  F.  and  Price,  J.  C.  C.,  Ala.  Agr.  Exp.  Sta.  Bui.  156  (1911). 

37.  Yule,  G.  U.,  Introduction  to  the  Theory  of  Statistics,  p.  146,  London 

(1919). 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  41 


Plate  I. — Blossom  buds  of  apple  on  February  2,  1920. 


Oldenburg 

Gano 

Daru 


Fameuse 

York 

Cilligos 


42 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


Plate  II. — By  rows:  left  to  right,  Oldenburg,  Primate,  Wealthy;  top  to 
bottom,  November  2,  1921,  January  28,  1922,  February  20,  1922. 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  43 


Plate  III. — By  rows:  left  to  right,  Rome,  Daru,  Cilligos;  top  to  bottom, 
November  2,  1921,  January  28,  1922,  February  20,  1922. 


44 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


Plate  IV. — Buds  developing  out  of  doors,  1920.  Left  to  right:  Cilligos, 

Fameuse,  Oldenburg. 


Plate  V. — Buds  forced  in  greenhouse,  photographed  March  6.  1922.  Cilligos 
(1),  Rome  (2)  (3),  Daru  (4),  Oldenburg  (5),  Primate  (6),  Fameuse  (7). 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  45 


APPENDIX 

Methods. — In  the  calculations  reported  in  this  paper,  the  date  of  first 
blossoming  has  been  used.  Though  this  standard  is  open  to  some  objections, 
as,  for  example,  a probable  fluctuation  with  the  total  quantity  of  blossoms  in 
the  tree,  it  is  less  subject  to  change  through  varying  judgments  of  different 
observers  than  is  the  date  of  full  bloom. 

Several  commentators  have  mentioned  the  variability  in  blossoming  found 
in  young  trees.  This  may  be  explained  by  the  fact  that  often  in  young  trees 
all  the  blossoms  are  on  terminal  shoots,  which  open  markedly  later  than  blos- 
soms on  spurs  and  if  recorded  without  qualifications  may  well  cause  a con- 
siderable change  in  the  relative  order  of  blossoming.  Phenological  records 
in  the  apple  should  distinguish  clearly  between  blossoms  on  spurs  and  those 
on  shoots.  All  samples  used  in  microscopic  study  were  gathered  from  spurs 
which  had  blossomed  at  least  once. 

Since  progress  in  phenological  studies  depends  on  the  availability  of  data, 
temperature  and  phenological  records  for  Columbia,  Missouri,  are  appended. 

DATES  OF  FIRST  BLOSSOMING  IN  APPLE  AT  COLUMBIA,  MO. 

(Days  of  the  year) 


Variety  1905  1906  1907  1908  1909  1911  1912  1913  1914  1915  1916  1917  1918 


Alexander  

114 

114 

94 

107 

115 

109 

116 

114 

113 

113 

109 

__ 

Arkansas 

98 

111 

86 

101 

108 

104 

113 

108 

111 

111 

106 

112 

101 

Arkansas  Beauty  

99 

110 

86 

100 

108 

101 

116 

109 

111 

111 

108 

108 

102 

Arkansas  Black  

101 

113 

88 

102 

116 

107 

115 

109 

111 

112 

107 

111 

104 

Ashton  _ _ 

97 

111 

86 

102 

108 

112 

118 

112 

112 

112 

109 





Autumn  Streaked  __ 

99 

110 



99 

109 

106 



107 

112 

111 

106 

109 

97 

Bailey  Sweet  

97 

110 

84 

101 

107 

101 

115 

107 

111 

109 

105 

108 

104 

Balagh  _ _ 

100 

111 

86 

100 

— 

103 





112 

112 

109 

113 

104 

Baldwin  

100 

111 

, 

103 

114 

107 

116 

110 

112 

112 

109 



96 

Battyani 



113 

93 

98 



103 

116 

110 

112 



108 



107 

Batullen  _ 

98 

92 



98 

113 

102 

116 

109 

112 

112 

108 

114 

101 

Ben  Davis  

99 

111 

85 

101 

113 

102 

115 

109 

111 

111 

108 

110 

102 

Ben  Hur  





87 

104 

113 

107 

116 

110 

112 

110 

107 

110 

96 

Black  Ben  Davis  __ 

98 

111 



103 



101 

119 

109 

112 

106 

Blenheim  _ _ _ 

108 

114 

95 

104 

116 

106 

118 

112 

113 

113 



114 

108 

Bosnian  

105 

112 

89 

103 



109 





113 



110 



105 

P.ripr 

95 

110 

87 

99 

109 

112 

112 

106 

113 

96 

Canada  Reinette 

99 

111 

87 

101 

109 

108 

115 

112 

Champion  

100 

110 

87 

103 



105 

118 

110 

112 

111 

109 



104 

Cilligos  — 

117 

120 

91 

113 



118 

115 

121 

117 

116 

117 



— 

Clark  _ _ _ _ — _ 

99 

111 



101 

113 

106 

116 

118 

113 



109 





Clayton  _ 

99 

111 

88 

102 

112 

104 

116 

110 

112 



108 





Collins  

102 

110 

86 

101 

109 

102 

114 

109 

111 

110 

107 

111 

107 

Czar  Thorn  

99 

111 

85 

99 

108 

104 

116 

109 



111 

109 

109 



Daru  

114 

116 

105 

112 

124 

112 

121 

113 

107 



114 



121 

Delaware  Red 



116 

88 

104 



_ 

115 

112 

115 

111 





105 

Delicious 



113 

87 

104 

113 

104 

116 

113 

112 

111 

107 

113 

102 

Devonshire  Duke  — 



112 

87 

102 



106 

118 

110 

116 

111 



113 

— 

Doctor  

101 

113 

90 

101 

115 

105 

115 

112 

112 

111 

109 

111 

106 

Downing  Blush  



113 

87 

102 



108 

118 

109 

110 

Eper  - 

115 

116 

99 

103 



112 



113 

114 

113 

114 

113 

108 

Fameuse  

96 

110 

86 

98 

108 

104 

115 

107 

109 

111 

107 

108 

91 

46 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


DATES  OF  FIRST  BLOSSOMING  IN  APPLE  AT  COLUMBIA,  MO. 

(Days  of  the  year) 


Variety  1905  1906  1907  1908  1909  1911  1912  1913  1914  1915  1916  1917  1918 


Faust’s  Rome  Beauty  113  115  99  107  119  105  — 114  113  113  109  __  __ 

Gano  98  111  86  102  113  103  116  110  111  112  109  114  101 

Ginnie  103  111  88  104  114  106  118  110  112  __  108  111  96 

Gold  Medal  100  114  87  102  113  105  119  111  115  112  109  __  102 

Golden  Russet  95  120  85  10'S  __  108  116  — 109  111  106  109  92 

Greening  102  111  87  102  116  106  117  111  — — __  __  __ 

Grimes  98  112  87  101  108  106  115  109  110  112  107  111  97 

Heidorn  __  115  __  107  116  106  115  111  112  112  112  113  105 

Hubbardston  99  111  86  103  114  100  115  112  114  111  108  113  97 

Huntsman  101  114  86  100  114  107  115  109  110  113  107  112  101 

Imperial  Janet 117  117  106  110  __  119  121  119  118  __  115  __  — 

Ingram  115  120  104  109  118  118  121  119  119  114  118  128  111 

Jeffries  __  __  — — __  106  116  110  112  111  107  113  96 

Jonathan  97  110  86  102  112  102  116  107  111  110  108  109  97 

July  100  112  87  100  „ 111  116  111  114  112  109  110  106 

Kansas  Greening  __  110  115  108  107  __  111  118  111  112  112  __  __  108 

Kartacs  __  116  108  106  118  __  __  115  115  113  110  114  104 

King  David  __  __  __  — 117  104  115  110  112  111  109  112  103 

Lady  116  115  99  108  __  „ — — — 111  114  121  __ 

Lady  Carter  99  110  87  101  113  __  — — 110  __  108  113  102 

London  __  115  88  104  112  __  __  __  117  112  109  — — 

Late  Duchess  97  109  87  98  107  __  __  __  109  112  106  121  92 

Longfield  100  111  95  100  109  — — — 115  __  112  __  106 

Lou  98  109  87  100  105  __  __  __  __  108  __  108  — 

Louise  100  111  86  101  111  __  __  __  112  112  109  112  96 

Magyar  105  111  88  103  — __  — 110  112  112  109  114  106 

Maiden  Blush  99  110  87  98  108  107  115  108  110  110  107  109  95 

Marin  97  110  86  100  109  104  115  107  109  111  105  109  97 

Melon  97  110  88  99  108  105  116  108  — — — — — 

Menagera  97  109  86  100  108  101  114  109  108  110  108  109  91 

Metitt  100  112  84  102  110  108  115  111  112  111  __  109  96 

Miller  Boy’s  Favorite  104  112  87  102  115  109  115  110  110  — __  „ — 

Minkler  98  110  85  99  107  102  119  107  108  109  107  108  96 

Minnesota  - 91  107  85  96  103  100  114  107  108  110  105  108  90 

Missing  Link  __  __  87  104  110  104  116  108  109  — 107  __  — 

Missouri  98  110  85  102  112  103  116  109  111  109  106  112  97 

Mosher  110  112  87  104  110  106  118  110  112  112  109  110  — 

Me  Intosh  100  111  87  102  109  105  117  110  112  111  109  113  __ 

Nelson  Sweet  100  112  87  103  114  106  116  110  112  113  __  — __ 

Noble  Savar  99  111  86  100  109  __  118  109  112  112  109  114  — 

Nyack  101  114  88  103  115  110  102  111  112  __  __  — — 

Nyari  Piros  105  112  86  99  109  104  115  110  109  114  107  __  100 

Ohio  Beauty  110  113  90  102  __  105  116  111  113  111  111  112  103 

Ohio  Pippin  99  110  85  100  llf2  103  116  108  113  111  108  109  97 

Oldenburg  94  110  86  98  107  102  114  107  110  __  106  — 97 

Olive  _ 99  112  87  102  111  106  — 111  112  113  109  114  __ 

Ontario  102  113  86  103  117  108  118  112  113  112  __  __  — 

Opalescent  106  116  110  117  114  113  111  — — 107 

Payne  Keeper  __  „ 93  105  114  106  117  113  112  113  109  114  98 

Peach  — 118  — 104  __  107  112  112  113  112  110  113  — 

Picket  100  110  86  99  110  105  __  __  110  112  109  109  96 

Ponyik  104  115  95  105  __  __  118  __  114  113  110  „ 106 

Primate  97  110  87  99  107  __  114  109  107  110  106  112  90 


Relation  of  Temperature  to  Blossoming — Apple  and  Peach  47 


DATES  OF  FIRST  BLOSSOMING  IN  APPLE  AT  COLUMBIA,  MO. 

(Days  of  the  year) 

Variety  1905  1906  1907  1908  1909  1911  1912  1913  1914  1915  1916  1917  1918 


Pumpkin  Russet  — 99  111  88  102  117  104  116  112  109  112  __  __  100 

Pumpkin  Sweet  100  111  87  101  108  __  __  „ 110  111  108  110  98 

Ralls  113  117  106  110  121  118  121  118  116  114  111  128  117 

Reagan  100  112  88  102  111  106  116  — — __  — — — 

Red  Astrachan  98  110  88  99  107  102  115  108  109  110  109  __  __ 

Red  June  __  _ __  __  — 104  111  __  112  111  109  113  94 

Red  Stettiner  99  110  85  100  112  103  116  108  113  111  108  109  97 

Rome  105  114  99  107  119  111  115  112  115  114  111  115  108 

Rutherford  96  109  85  99  — — 115  109  112  111  108  110  92 

Sabadka  100  115  87  __  — 106  118  112  113  113  109  110  103 

Segfu  — 112  87  104  113  110  117  112  112  112  110  110  102 

Sekula  104  113  94  102  __  110  __  __  115  __  109  — 102 

Selumes  100  112  85  102  107  107  115  114  111  112  108  — 98 

Skelton  99  111  87  102  112  __  __  110  114  112  108  __  104 

Spitzenberg  99  111  87  102  109  __  __  111  117  „ __  __  __ 

Standard  93  110  86  99  104  100  __  108  110  112  106  — 92 

Stayman  98  110  86  101  112  106  115  110  111  111  109  112  103 

Summer  Calville  __  97  110  86  98  108  105  116  109  109  — 108  110  92 

Summer  King  — __  __  — __  114  113  — 113  109  112  103 

Tetofski  98  110  87  103  108  106  115  109  112  110  106  110  100 

Titus  Pippin  102  110  86  102  114  106  115  109  111  110  108  111  105 

Tudor  99  115  87  — __  110  116  109  109  __  106  __  98 

Wafer  94  112  88  102  113  114  __  110  114  112  110  112  106 

Wealthy  __  __  __  104  112  104  117  109  112  111  109  110  103 

White  Canada  98  112  87  101  109  105  117  108  111  112  108  112  97 

White  Pippin  97  112  87  102  __  107  116  — 112  __  106  — 94 

Wine  Rubets  104  112  87  104  117  109  117  110  109  112  109  — 102 

Winesap  99  111  87  102  113  104  116  110  112  111  109  109  104 

Wolf  River  108  113  88  102  112  109  116  110  111  111  110  113  102 

Woodmansee  ~ — — — — 118  113  114  112  110  114  121 

Workaroe  109  113  — 103  115  109  118  112  115  111  __  __  — 

Yappa  102  112  87  102  116  106  118  108  — __  __  __  — 

Yellow  Newtown __  __  __  104  __  108  117  113  113  112  109  __  103 

Yellow  Transparent  — — — 108  113  104  117  113  113  112  108  110  105 

York  Imperial  99  113  88  103  113  106  116  111  — 112  __  112  __ 

York  Stripe  103  114  88  104  116  107  117  113  — __  — __  __ 


48 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


DAILY  MAXIMUM  TEMPERATURES,  COLUMBIA,  MISSOURI 

January 


1905 

1906 

1907 

1908 

1909 

1910 

1911 

1912 

1913 

1914 

1915 

1916 

1917 

1918 

1 

66 

39 

40 

47 

28 

46 

52 

26 

56 

42 

41 

62 

43 

44 

2 

43 

41 

49 

50 

49 

29 

7 

26 

38 

42 

33 

42 

50 

43 

3 

27 

41 

47 

49 

59 

23 

5 

19 

42 

29 

38 

48 

49 

31 

4 

35 

29 

46 

47 

60 

29 

34 

11 

36 

29 

41 

61 

50 

44 

5 

40 

42 

65 

42 

33 

25 

50 

13 

23 

28 

47 

63 

42 

42 

6 

25 

41 

72 

51 

4 

12 

48 

—3 

20 

39 

42 

21 

56 

28 

7 

24 

43 

71 

52 

17 

35 

55 

2 

20 

54 

43 

29 

40 

21 

8 

32 

19 

54 

44 

33 

36 

47 

12 

24 

58 

30 

38 

54 

26 

9 

31 

40 

26 

52 

43 

29 

47 

29 

32 

40 

49 

49 

57 

20 

10 

10 

45 

33 

47 

42 

43 

68 

10 

38 

28 

40 

51 

52 

16 

11 

32 

35 

44 

36 

8 

42 

67 

3 

38 

44 

40 

37 

22 

9 

12 

15 

35 

43 

30 

18 

44 

31 

—5 

20 

34 

41 

34 

37 

0 

13 

11 

43 

56 

31 

32 

44 

34 

20 

32 

46 

52 

0 

16 

15 

14 

9 

48 

40 

39 

35 

30 

34 

22 

43 

52 

54 

20 

30 

18 

15 

17 

58 

28 

44 

34 

33 

22 

7 

56 

58 

61 

34 

23 

16 

16 

30 

37 

33 

26 

30 

37 

27 

29 

60 

53 

58 

11 

22 

23 

17 

39 

57 

34 

40 

27 

51 

28 

43 

59 

43 

28 

18 

30 

18 

18 

40 

40 

54 

43 

36 

44 

29 

33 

38 

50 

31 

23 

30 

13 

19 

37 

63 

60 

49 

37 

57 

41 

20 

64 

66 

34 

32 

40 

17 

20 

46 

72 

26 

58 

46 

44 

54 

27 

35 

47 

25 

55 

40 

16 

21 

34 

65 

49 

58 

61 

32 

39 

48 

32 

32 

20 

57 

60 

20 

22 

24 

17 

39 

44 

74 

42 

33 

49 

37 

54 

19 

52 

17 

24 

23 

33 

24 

35 

35 

72 

50 

45 

51 

42 

60 

14 

54 

36 

37 

24 

31 

43 

51 

32 

65 

44 

49 

34 

41 

41 

16 

62 

34 

42 

25 

10 

47 

23 

47 

40 

62 

55 

39 

54 

38 

26 

60 

45 

42 

26 

30 

48 

18 

42 

43 

61 

71 

50 

53 

62 

31 

63 

41 

28 

27 

37 

48 

25 

42 

57 

41 

60 

35 

43 

64 

26 

56 

42 

15 

28 

29 

47 

32 

42 

61 

42 

47 

34 

43 

65 

15 

33 

66 

18 

30 

25 

48 

33 

31 

10 

29 

40 

28 

54 

30 

42 

31 

51 

19 

31 

27 

46 

35 

34 

18 

32 

64 

41 

31 

43 

51 

27 

47 

5 

Relation  of  Temperature  to  Blossoming — Apple  and  Peach  49 


DAILY  MAXIMUM  TEMPERATURES,  COLUMBIA,  MISSOURI 

February 


1905 

1906 

1907 

1908 

1909 

1910 

1911 

1912 

1913 

1914 

1915 

1916 

1917 

1918 

1 

15 

44 

37 

17 

43 

57 

81 

33 

14 

52 

48 

16 

6 

11 

2 

—2 

27 

36 

31 

55 

51 

38 

21 

26 

55 

32 

17 

3 

30 

3 

5 

55 

14 

35 

63 

38 

54 

20 

32 

38 

38 

21 

36 

24 

4 

20 

42 

11 

37 

65 

43 

41 

18 

30 

32 

56 

36 

38 

11 

5 

20 

14 

12 

52 

60 

44 

46 

17 

15 

35 

37 

36 

24 

53 

6 

21 

21 

21 

36 

45 

31 

43 

23 

27 

32 

31 

22 

48 

50 

7 

23 

30 

25 

49 

43 

40 

41 

24 

25 

15 

34 

14 

47 

59 

8 

28 

40 

47 

41 

54 

41 

46 

22 

34 

16 

38 

35 

38 

65 

9 

29 

27 

50 

43 

53 

32 

40 

20 

35 

45 

45 

41 

24 

40 

10 

13 

29 

41 

46 

33 

43 

47 

34 

40 

37 

65 

44 

29 

63 

11 

27 

43 

49 

46 

56 

42 

48 

39 

36 

28 

64 

44 

21 

65 

12 

9 

58 

57 

62 

54 

25 

54 

30 

21 

21 

69 

35 

33 

54 

13 

—1 

51 

60 

54 

35 

42 

64 

37 

38 

22 

61 

21 

44 

59 

14 

24 

28 

43 

40 

36 

59 

54 

36 

52 

24 

49 

33 

35 

62 

is 

15 

28 

60 

38 

21 

63 

78 

41 

51 

38 

34 

43 

38 

30 

16 

30 

37 

60 

34 

25 

17 

76 

53 

61 

27 

51 

56 

56 

32 

17 

30 

36 

62 

39 

49 

17 

65 

53 

63 

54 

57 

52 

52 

36 

18 

35 

55 

66 

37 

46 

30 

39 

55 

69 

42 

55 

40 

33 

51 

19 

35 

61 

44 

23 

38 

39 

32 

45 

63 

27 

51 

55 

52 

60 

20 

34 

59 

43 

37 

54 

41 

26 

35 

38 

30 

54 

55 

33 

16 

21 

44 

57 

25 

34 

58 

26 

22 

30 

51 

48 

52 

59 

53 

19 

22 

47 

69 

26 

56 

54 

26 

31 

40 

34 

38 

50 

63 

64 

45 

23 

55 

54 

31 

49 

55 

10 

35 

52 

23 

17 

47 

37 

56 

69 

24 

51 

50 

35 

49 

35 

28 

43 

43 

26 

21 

34 

49 

37 

65 

25 

50 

45 

50 

48 

55 

43 

48 

39 

40 

32 

41 

43 

68 

63 

26 

60 

39 

54 

36 

56 

42 

39 

35 

40 

46 

35 

37 

62 

51 

27 

44 

32 

58 

31 

52 

33 

34 

38 

29 

53 

35 

32 

31 

42 

28 

63 

50 

61 

52 

65 

48 

29 

33 

22 

50 

40 

31 

41 

38 

29 

.... 

71 

.... 

.... 

27 

.... 

.... 

.... 

38 

.... 

50 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  53 


DAILY  MAXIMUM  TEMPERATURES,  COLUMBIA,  MISSOURI 

March 


1905 

1906 

1907 

1908 

1909 

1910 

1911 

1912 

1913 

1914 

1915 

1916 

1917 

1918 

1 

59 

62 

54 

48 

67 

57 

34 

31 

28 

27 

44 

32 

36 

50 

2 

67 

59 

59 

48 

70 

70 

46 

24 

31 

39 

42 

32 

34 

58 

3 

76 

34 

54 

45 

49 

76 

59 

29 

49 

35 

42 

26 

31 

51 

4 

58 

33 

40 

49 

48 

76 

46 

33 

41 

43 

31 

50 

22 

62 

5 

67 

34 

51 

74 

62 

81 

70 

27 

38 

50 

33 

53 

38 

76 

6 

39 

40 

41 

71 

54 

64 

59 

37 

30 

40 

29 

66 

57 

42 

7 

44 

42 

49 

46 

62 

58 

49 

36 

52 

34 

34 

43 

50 

53 

8 

50 

60 

45 

40 

46 

57 

63 

33 

67 

37 

39 

38 

50 

65 

9 

53 

52 

38 

47 

42 

42 

71 

28 

57 

47 

41 

56 

64 

73 

10 

43 

39 

39 

56 

43 

48 

61 

31 

55 

48 

39 

52 

69 

47 

11 

43 

32 

53 

50 

39 

60 

82 

35 

59 

34 

37 

49 

54 

65 

12 

50 

22 

55 

73 

46 

61 

60 

36 

61 

47 

41 

76 

44 

77 

13 

47 

29 

37 

55 

51 

69 

49 

40 

57 

62 

45 

76 

54 

90 

14 

69 

25 

43 

74 

37 

47 

60 

37 

54 

69 

49 

51 

45 

61 

15 

70 

25 

63 

66 

45 

52 

50 

33 

28 

79 

39 

36 

54 

50 

16 

71 

23 

66 

66 

48 

65 

45 

51 

39 

55 

46 

52 

60 

56 

17 

66 

26 

68 

63 

52 

67 

64 

58 

63 

53 

44 

56 

41 

73 

18 

71 

29 

67 

63 

72 

71 

52 

65 

68 

36 

42 

62 

41 

76 

19 

55 

30 

80 

45 

48 

74 

60 

67 

68 

30 

42 

54 

65 

72 

20 

36 

34 

70 

51 

48 

70 

72 

41 

50 

37 

33 

71 

49 

74 

21 

48 

46 

92 

61 

50 

73 

77 

28 

34 

37 

37 

86 

64 

80 

22 

73 

32 

90 

60 

60 

90 

59 

32 

45 

46 

37 

64 

71 

62 

23 

61 

29 

82 

67 

69 

88 

53 

34 

72 

58 

47 

54 

64 

42 

24 

71 

35 

77 

67 

66 

85 

58 

39 

53 

68 

63 

82 

66 

59 

25 

71 

55 

90 

81 

45 

82 

71 

48 

39 

70 

50 

69 

75 

69 

26 

71 

59 

82 

73 

67 

86 

61 

49 

33 

73 

39 

56 

59 

75 

27 

82 

42 

82 

78 

49 

86 

50 

42 

32 

61 

48 

51 

50 

64 

28 

69 

39 

78 

44 

55 

86 

66 

43 

53 

77 

53 

63 

69 

61 

29 

60 

37 

58 

57 

45 

78 

55 

59 

62 

67 

46 

62 

62 

66 

30 

73 

53 

61 

48 

52 

65 

46 

68 

75 

58 

44 

68 

82 

72 

31 

77 

49 

47 

67 

49 

60 

45 

72 

63 

69 

40 

59 

80 

75 

Relation  of  Temperature  to  Blossoming — Apple  and  Peach  51 


DAILY  MAXIMUM  TEMPERATURES,  COLUMBIA,  MISSOURI 

April 


1905 

1906 

1907 

1908 

1909 

1910 

1911 

1912 

1913 

1914 

1915 

1916 

1917 

1918 

1 

81 

55 

54 

57 

57 

71 

48 

50 

72 

57 

40 

52 

50 

79 

2 

72 

70 

63 

38 

56 

69 

69 

57 

79 

63 

46 

50 

54 

83 

3 

70 

74 

63 

60 

61 

74 

43 

65 

72 

46 

54 

61 

62 

51 

4 

58 

61 

70 

65 

85 

76 

47 

74 

50 

52 

71 

59 

54 

55 

5 

55 

56 

50 

65 

84 

56 

55 

80 

65 

60 

75 

56 

54 

54 

6 

55 

64 

50 

80 

76 

60 

63 

72 

69 

56 

76 

45 

59 

58 

7 

61 

73 

57 

69 

59 

73 

49 

55 

53 

46 

75 

36 

51 

60 

8 

86 

69 

47 

70 

48 

77 

57 

66 

46 

36 

77 

44 

46 

53 

9 

90 

72 

52 

54 

51 

75 

58 

67 

65 

46 

70 

50 

57 

48 

10 

74 

66 

51 

63 

59 

81 

62 

68 

48 

53 

69 

68 

70 

54 

11 

59 

78 

54 

59 

73 

63 

54 

77 

44 

49 

63 

82 

70 

53 

12 

62 

82 

47 

69 

61 

58 

78 

76 

43 

57 

58 

84 

57 

52 

13 

64 

66 

45 

79 

55 

72 

66 

74 

57 

62 

60 

70 

51 

64 

14 

53 

45 

52 

68 

64 

71 

53 

74 

68 

65 

71 

59 

59 

66 

15 

48 

54 

62 

68 

65 

68 

62 

65 

72 

72 

77 

68 

52 

61 

16 

46 

63 

45 

57 

80 

47 

70 

49 

77 

83 

81 

62 

73 

73 

17 

55 

69 

51 

61 

87 

42 

77 

44 

84 

83 

79 

65 

83 

65 

18 

61 

74 

44 

64 

78 

40 

61 

55 

79 

70 

76 

79 

83 

67 

19 

66 

77 

47 

81 

48 

42 

62 

53 

66 

51 

84 

76 

80 

55 

20 

75 

75 

56 

83 

51 

66 

61 

70 

63 

64 

78 

70 

62 

38 

21 

61 

81 

60 

80 

55 

78 

61 

73 

81 

84 

76 

55 

72 

50 

22 

64 

64 

65 

82 

54 

65 

58 

58 

82 

81 

78 

67 

81 

63 

23 

70 

61 

70 

79 

60 

40 

59 

68 

80 

67 

86 

65 

86 

63 

24 

73 

87 

78 

73 

69 

36 

60 

75 

60 

81 

84 

61 

81 

48 

25 

73 

80 

72 

75 

71 

42 

63 

65 

60 

81 

84 

57 

61 

45 

26 

59 

82 

57 

55 

82 

59 

75 

72 

64 

86 

78 

51 

53 

58 

27 

80 

86 

65 

48 

64 

77 

58 

67 

57 

81 

83 

61 

49 

58 

28 

85 

70 

81 

54 

80 

85 

76 

66 

66 

71 

85 

69 

54 

62 

29 

72 

77 

72 

52 

88 

91 

82 

57 

73 

58 

73 

73 

49 

55 

30 

73 

61 

47 

57 

51 

80 

72 

67 

84 

53 

73 

64 

54 

56 

UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 


AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  54 


Studies  In  Animal  Nutrition 

II.  Changes  in  Proportions  of  Carcass 
and  Offal  on  Different  Planes 
of  Nutrition 


(Publication  authorized  September  1,  1922.) 


COLUMBIA,  MISSOURI 
SEPTEMBER,  1922 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL 

the;  curators  of  the  university  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY  P.  E.  BURTON  H.  J.  BLANTON 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 

J.  C.  JONES,  PH.  D.,  LL.  D.,  PRESIDENT  OF  THE  UNIVERSITY 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 


STATION  STAFF 

SEPTEMBER.  1922 


AGRICULTURAL  CHEMISTRY 

C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  Ph.  D. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  SiEveking,  B.  S.  in  Agr. 

AGRICULTURAL  ENGINEERING 
J.  C.  Wooley,  B.  S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

DAIRY  HUSBANDRY 
A.  C.  Ragsdale,  B.  S.  in  Agr. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

W.  P.  Hays 


ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride,  B.  S.  in  Agr. 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  Ph.  D. 

O.  W.  Letson,  B.  S.  in  Agr. 
Miss  Regina  Schulte* 


RURAL  LIFE 
O.  R.  Johnson,  A.  M. 

S.  D.  Gromer,  A.  M. 

E.  L.  Morgan,  A.M. 

Ben  H.  Frame,  B.  S.  in  Agr. 
Owen  Howells,  B.  S.  in  Agr. 


HORTICULTURE 
T.  J.  Talbert,  A.  M. 

H.  D.  Hooker,  Jr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  Ph.  D. 

H.  G.  Swartwout,  B.  S.  in  Agr. 

J.  T.  Quinn,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY 
H.  L.  Kempster,  B.  S. 

Earl  W.  Henderson,  B.S. 

SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M 
W.  A.  Albrecht,  Ph.  D. 

F.  L.  Duley,  A.M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr. 

Richard  Bradfield,  Ph.  D. 

VETERINARY  SCIENCE 
J.  W.  Connaway,  D.  V.  S.,  M.  D. 

L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 

OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Secretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Jane  Frodsham,  Librarian. 

E.  E.  Brown,  Business  Manager. 


In  service  of  U.  S.  Department  of  Agriculture. 


STUDIES  IN  ANIMAL  NUTRITION 

II.  Changes  in  Proportions  of  Carcass  and  Offal  on 
Different  Planes  of  Nutrition. 

C.  Robert  Moulton,  P.  F.  Trowbridge,*  L.  D.  Haigh 

The  changes  experienced  by  beef  cattle  in  form  and  weight 
when  on  different  planes  of  nutrition  were  presented  and  discussed 
in  a previous  bulletinf.  Representative  animals  from  each  of  the 
groups  were  killed  at  intervals  from  birth  to  four  years  old.  The 
data  collected  in  the  slaughter  house  will  be  presented  in  this  bul- 
letin. 


RATION 

For  a general  discussion  of  the  treatment  of  the  animals  the 
previous  bulletin  must  be  consulted.  The  ration  included  milk  for 
several  months  after  birth  and  timothy  hay  and  grain  were  soon 
introduced.  At  weaning  time  the  ration  consisted  of  alfalfa  hay 
and  a grain  mixture  in  the  ratio  of  one  to  two.  The  grain  consisted 
of  six  parts  corn  chop,  three  parts  whole  oats,  and  one  part  of  old 
process  linseed  meal. 

PLANE  OF  NUTRITION 

The  animals  were  early  divided  into  three  groups.  Group  I 
was  fed  all  it  would  eat  of  the  ration.  Group  II  was  fed  for  maxi- 
mum growth  without  permitting  the  laying  on  of  much  fat.  Group 
III  was  fed  for  scanty  or  retarded  growth.  The  Group  II  steers 
gained  about  a pound  a day  for  the  first  two  years  while  the  Group 
III  cattle  gained  but  0.69  pounds  per  day. 

SLAUGHTERING 

In  the  conduct  of  this  experiment  great  care  was  exercised  that 
all  of  the  operations  with  the  different  animals  should  be  carried 
out  under  similar  conditions.  Although  the  slaughtering  and  sub- 
sequent operations  were  necessarily  carried  out  at  different  times, 


•Resigned  September,  1918. 

tC.  Robert  Moulton,  P.  F.  Trowbridge,  L.  D.  Haigh,  Studies  in  Animal  Nutrition,  I. 
Changes  in  Form  and  Weight  on  Different  Planes  of  Nutrition,  Mo.  Agr.  Expt.  Station, 
Research  Bulletin  43. 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


in  order  to  make  the  results  strictly  comparable  the  slaughtering 
and  cutting  were  done  by  the  same  experts  each  time*. 

Each  animal  was  slaughtered  on  the  day  following  the  closing 
of  its  feeding  period.  In  the  morning  the  steer  was  fed  and  weighed 
as  usual  but  no  water  was  given.  If  slaughter  occurred  late  in  the 
morning  or  in  the  afternoon  the  animal  was  weighed  again  im- 
mediately before  slaughtering.  The  animal  was  led  from  the  feed- 
ing shed  to  the  slaughter  house  followed  by  men  with  a shovel  and 
long  handled  dipper  to  catch  any  feces  or  urine  that  might  be 
voided. 

The  animal  was  stunned  with  a knocking  hammer,  shackled 
by  the  hind  legs  and  hoisted  to  swing  clear  of  the  floor.  An  oil 
cloth,  funnel  shaped  bag  had  been  fastened  to  the  muzzel  before 
hoisting  so  that  any  vomit  voided  could  be  caught  and  weighed. 
The  suspended  animal  was  stuck  in  the  throat  near  the  brisket  so 
that  both  the  jugular  vein  and  carotid  artery  were  severed.  Com- 
plete bleeding  was  assured  by  pumping  the  fore  legs  up  and  down. 
The  blood  was  caught  in  a tared  pan  and  weighed.  The  volume 
of  this  main  weighed  portion  was  determined.  A tared  pan  was 
kept  under  the  animal  to  catch  any  blood  that  might  drip  while 
skinning  out  the  head. 

While  the  blood  was  flowing  freely  the  samples  for  analysis 
were  taken  and  poured  in  suitable  quantity  into  tared,  covered 
containers  and  crucibles.  The  blood  was  still  warm  and  no  clotting 
had  yet  occurred. 

While  the  carcass  was  hanging  the  head  was  skinned  out.  The 
gullet  was  firmly  tied  with  a skewer  thrust  through  below  the  tie 
and  the  head  was  then  severed  permitting  the  dripping  blood  to 
collect  in  a tared  container.  The  head  was  immediately  placed  in 
a large  can  provided  with  a cover  and  the  weight  obtained.  The 
tongue  with  larynx  and  bones  was  removed,  separated  into  tongue 
marketable,  tongue  base,  bones,  larynx,  and  piece  of  gullet.  The 
parts  were  weighed  and  set  aside  in  closed  containers.  Horns  were 
sawed  off  and  weighed.  The  skull  and  entire  head  was  split  accur- 
ately in  half  and  the  brain  was  removed  and  weighed.  The  lean 
meat  and  fat  were  removed  from  the  right  half  as  were  the  teeth. 
If  any  vomit  was  found  it  was  weighed  and  discarded.  Both  halves 
of  the  head  were  weighed.  The  total  lean  and  fat  were  obtained 


* Swift  & Company  of  Kansas  City  furnished  an  expert  butcher  to  do  the  slaughtering. 
Mr.  Samuel  Godfrey,  foreman  of  the  beef  cutting  department,  did  the  cutting. 


Studies  In  Animal  Nutrition — II 


5 


by  doubling  the  right-side  weights  and  the  total  bone  by  subtract- 
ing the  sum  of  the  lean  and  fat  from  the  sum  of  the  two  halves  of 
the  head  minus  the  teeth. 

As  soon  as  the  suspended  carcass  with  the  head  removed  had 
practically  stopped  bleeding  it  was  lowered  to  the  floor  with  the 
anterior  end  lying  up  the  slope  of  the  smooth  cement  floor.  A man 
with  a rubber  window  wiper  and  sharp  edged  dust  pan  kept  all 
oozing  blood  wiped  up  and  transferred  to  weighed  containers. 

In  skinning  out  the  feet  the  dew  claws  were  removed,  weighed 
and  saved  and  care  was  taken  to  remove  the  hide  exactly  at  the 
hoof  line.  From  the  right  feet  the  hoofs  were  separated  and  the 
remainder  of  the  material  was  considered  as  skeleton.  The  usual 
packing  house  order  of  procedure  was  followed  in  skinning  the 
carcass  and  removing  the  internal  organs.  The  caul  fat  was  re- 
moved while  the  carcass  was  on  its  back.  The  bladder  was  tied 
before  removing  and  weighed  with  its  contents  and  again  empty. 
The  rectum  was  tied  as  soon  as  it  was  cut  loose.  The  tail  was 
removed  and  split  in  two  and  the  lean  and  fat  meat  separated  and 
weighed. 

The  contents  of  the  abdominal  and  thoracic  cavities  was  caught 
in  a large  tub,  or  tubs,  and  weighed.  The  separation  and  weighing 
of  the  organs  was  pushed  as  rapidly  as  possible  to  reduce  to  a 
minimum  the  loss  of  water  by  evaporation.  A double  tie  was  made 
at  the  end  of  the  small  intestine  near  the  abomasum  before  severing 
one  from  the  other.  The  fat  was  carefully  cut  or  scraped  from  all 
four  stomachs  and  weighed.  The  intestines  were  also  carefully 
freed  from  fat,  and  this  was  generally  accomplished  without  any 
portion  becoming  smeared  with  the  contents.  The  stomachs  were 
emptied  of  the  contents  and  were  cleaned  by  washing  with  water 
after  which  they  were  wiped  dry  with  cloths.  The  contents  of  the 
intestines  were  removed  by  stripping  through  the  fingers  taking 
a section  at  a time.  The  stripped  sections  were  split  open  and  the 
inside  was  wiped  lightly  to  remove  all  contents.  Occasionally  it 
was  necessary  to  wash  and  wipe  a smeared  portion.  The  various 
organs  were  separated,  weighed  and  put  into  closed  containers. 

The  hide  was  removed  and  the  carcass  was  split  into  halves. 
Then  the  spinal  cord  was  removed.  The  diaphragm  was  removed 
back  to  the  striated  muscle  and  composited  with  the  internal  or- 
gans. 

The  carcass  was  allowed  to  chill  for  48  hours  and  the  right 


6 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


side  was  separated  into  the  two  quarters  and  then  into  the  standard 
wholesale  cuts.  (Figure  1).  Each  cut  was  separated  into  lean  meat, 
fatty  tissue  and  bone.  The  tendons  were  weighed  with  the  bone. 
In  the  separation  of  the  lean  and  fat,  care  was  taken  that  the  fatty 
tissue  should  contain  no  lean  meat.  Necessarily  the  lean  meat 
contained  small  pieces  of  fat  which  could  not  be  separated.  The 
meat  was  cut  from  the  bones  as  completely  as  could  be  done  with 
a boning  knife.  All  samples  were  kept  in  closed  tin  containers. 

The  further  treatment  of  the  separated  parts  is  of  interest  only 
in  connection  with  the  chemical  analysis  and  the  description  will  be 
deferred  to  a later  publication. 

THE  SLAUGHTER  HOUSE  DATA 

The  detailed  slaughter  house  data  obtained  are  given  in  the 
Appendix  in  Tables  1 to  5 for  the  offal  parts  and  Tables  6 to  10  for 
the  carcass  parts.  Few  of  the  weights  need  explanation.  The 
warm  empty  weight  was  obtained  by  subtracting  the  contents  of 
the  stomachs,  intestines,  and  urinary  bladder  and  any  excrement 
voided  before  death  from  the  live  weight  at  slaughtering.  The 
heart  and  neck  sweetbreads  are  the  thymus  gland.  The  stomach 
and  intestinal  fats  are  those  which  adhered  to  the  respective  organs. 
The  intestinal  fat  is  largely  included  in  the  mesentery.  The  caul 
fat  is  that  laid  on  in  the  part  of  the  peritoneum  stretching  like  an 
apron  over  the  stomachs  and  intestines.  The  different  divisions 
are  mutually  exclusive  excepting  where  specified  otherwise. 

Table  I. — Percent  Empty  Weight  to 
Live  Weight. 


Age 

Group 

Group  Group 

I 

II 

III 

At  birth  

. 98.41 

98.41 

98.41 

3 months  

. 88.02 

89.27 

83.89 

5%  months  

. 84.32 

85.29 

86.63 

8^  months  

. 83.16 

82.28 

83.19 

11  months  

. 88.30 

87.61 

87.10 

18  months  

. 88.77 

90.37 

21-26  months  

. 90.44 

88.69 

87.05 

34  months  

. 90.39 

91.63 

38  months  stunted  . 

. 92.30 

40  months  

. 93.26 

89.19 

88.95 

45  months  

. 90.40 

87.68 

89.01 

47-48  months  

. 92.24 

90.12 

89.09 

Empty  Weight. — Table  I gives  the  proportion  of  empty  weight 
in  the  live  animal  from  birth  to  four  years  for  each  group  of  cattle. 
The  figures  in  this  table  as  well  as  in  Tables  II  to  XVIII  inclusive 


Studies  In  Animal  Nutrition — II 


7 


are  taken  from  Tables  11  to  15  in  the  Appendix.  The  figures  pre- 
sented for  the  animal  at  birth  are  for  the  average  of  Hereford  calves 
reported  in  Research  Bulletin  38  of  this  Station.  The  comple- 
ment of  the  percent  of  empty  weight  is  of  course  the  percent  of 
fill. 

The  animal  at  birth  has  the  largest  percent  of  empty  weight. 
This  is  due  to  the  lack  of  food  and  food  residues  in  the  alimentary 
canal.  The  figure  decreases  during  the  early  months  showing  a 
large  proportion  of  fill  and  is  least  at  8^2  months.  It  then  gradually 
increases  with  some  irregularity  and  becomes  at  about  3 years  and 
thereafter  a higher  proportion  than  at  any  time  since  birth. 

The  amount  of  the  ration  afifects  the  percent  of  the  empty 
weight.  The  lighter  rations  show  generally  a smaller  percent  of 
empty  weight  and  consequently  indicate  more  relative  fill.  Since 
the  weight  of  the  ration  is  smaller  and  the  weight  of  fill  is  smaller 
this  can  only  be  due  to  a greater  difference  in  weight  of  animal  re- 
sulting in  a decreased  percent  of  empty  weight.  At  8 3/2  months 
there  is  practically  no  difference  between  the  three  groups. 

Carcass. — Tables  II  and  III  show  the  percent  of  carcass  in  the 
live  and  empty  animals  respectively.  The  percent  of  carcass  to  live 
weight  decreases  to  about  8*4  months  and  then  increases  to  reach 
at  3 to  4 years  a higher  value  than  at  any  previous  time.  For  the 
first  six  months  there  is  little  difference  between  the  groups  but 
thereafter  the  better  fed  animals  have  the  greater  percent  of  carcass. 

The  effect  of  varying  proportions  of  fill  is  shown  in  the  figures 
discussed  in  the  above  paragraph.  On  the  empty  weight  basis 
there  is  a continuous  increase  in  percent  of  carcass  from  birth  to 


Table  II. — Percent  Carcass  to  Live 
Weight. 


Table  III. — Percent  Carcass  to  Empty 
Weight. 


Age 

At  birth  

3 months  

5%  months  

8%  months  

11  months  

18  months  

21-26  months  

34  months  

38  months  stunted  . . 

40  months  

45  months  

47-48  months  


Group 

Group 

Group 

I 

II 

III 

59.30 

59.30 

59.30 

54.19 

57.32 

53.63 

53.71 

53.00 

54.39 

54.17 

50.46 

51.09 

57.52 

53.85 

55.23 

60.46 

56.02 

58.24 

57.71 

54.27 

61.49 

60.49 

65.69 

70.42 

61.18 

58.08 

65.32 

60.28 

60.34 

68.95 

61.80 

59.63 

Age 

At  birth  

3 months  

5%  months  

8%  months  

11  months  

18  months  

21-26  months  

34  months  

38  months  stunted  . . 

40  months  

45  months  

47-48  months  


Group 

Group 

Group 

I 

II 

III 

60.39 

60.39 

60.39 

61.56 

64.21 

63.92 

63.70 

62.14 

62.78 

65.14 

61.33 

61.41 

65.14 

61.47 

63.41 

68.11 

61.98 

64.39 

65.07 

62.34 

68.03 

66.02 

71.17 

75.52 

68.59 

65.29 

72.25 

68.76 

67.80 

74.75 

68.58 

66.63 

8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


3 or  4 years  of  age.  The  relation  between  the  groups  is  the  same 
as  when  the  live  weight  formed  the  basis. 

Carcass  and  Offal  Fat. — In  Tables  IV  and  V are  shown  the 
percents  of  carcass  plus  offal  fat  to  live  and  empty  weights.  The 
offal  fat  is  the  hand  separable  fat  on  the  internal  organs.  It  is 
small  in  amount  in  the  young  animals,  being  entirely  negligible  in 
the  calf  at  birth.  In  the  very  fat  four  year  old  steer  it  amounts  to 
nearly  five  percent  of  the  animal.  The  effect  of  adding  this  fat  to 
the  carcass  weight  is  merely  to  increase  the  spread  of  the  figures 
with  increasing  age  and  fatness.  The  general  relations  pointed  out 
in  the  section  above  hold  here. 


Table  IV. — Percent  Carcass  and  Oefal  Table  V. — Percent  Carcass  and  Oeeal 

Fat  to  Live  Weight.  Fat  to  Empty  Weight. 


Age 

Group 

Group  Group 

Age 

Group 

Group  Group 

I 

II 

III 

I 

II 

III 

At  birth  

. 59.30 

59.30 

59.30 

At  birth  

. 60.39 

60.39 

60.39 

3 months  

. 55.45 

58.06 

54.17 

3 months  

. 62.99 

65.03 

64.57 

5%  months  

. 57.01 

54.53 

54.23 

5%  months  

. 67.61 

63.94 

63.76 

8 y%  months  

. 56.05 

52.23 

51.85 

8%  months  

. 67.40 

63.48 

62.33 

11  months  

. 61.16 

56.17 

56.69 

1 1 months  

. 69.26 

64.12 

65.08 

18  months  

. 65.04 

57.40 

18  months  

. 73.27 

63.49 

21-26  months  

. 63.01 

59.79 

55.89 

21-26  months  

. 69.67 

67.41 

64.21 

34  months  

. 65.51 

62.96 

34  months  

. 72.47 

68.71 

38  months  stunted  . 

. 71.34 

38  months  stunted  . 

. 77.30 

40  months  

. 76.18 

63.58 

59.46 

40  months  

. 81.69 

71.29 

66.84 

45  months  

. 71.62 

62.53 

62.60 

45  months  

. 79.22 

71.32 

70.33 

47-48  months  

. 73.33 

64.98 

62.19 

47-48  months  

. 79.49 

72.11 

69.80 

Table  VI. — Percent  Offal  Fat  to  Empty 
Weight. 


Age 

Group 

Group  Group 

I 

II 

III 

At  birth  

none 

none 

3 months  

1.43 

0.83 

0.65 

5%  months  

..  3.91 

1.80 

0.98 

8%  months  

2.26 

2.16 

0.92 

1 1 months  

4.12 

2.65 

1.68 

18  months  

5.16 

1.51 

21-26  months  

5.28 

2.34 

1.87 

34  months  

4.45 

2.70 

38  months  stunted  . 

6.13 

40  months  

6.17 

2.70 

1.55 

46  months  

. 6.97 

2.56 

2.54 

47-48  months  

4.74 

3.53 

3.17 

Table  VI  gives  the  percent  of  offal  fat  to  the  empty  animal. 
There  is  a rather  consistent  increase  in  percent  of  offal  fat  with 
increasing  age  and  fatness.  A striking  exception  is  shown  by  the 
four  year  old  Group  I steer.  This  is  due  to  a great  decrease  in  the 


Fig.  1. — Wholesale  divisions  of  the  beef  carcass. 


Studies  In  Animal  Nutrition — II 


9 


weight  of  offal  fat,  this  animal  having  but  38.5  kilograms  while  the 
next  younger  animals  had  53.6  and  48.3  kilograms  respectively. 
The  low  figure  should  be  taken  with  reservations  as  to  its  general 
applicability. 

Hide  and  Hair. — The  percent  of  hide  and  hair  referred  to  empty 
weight  is  given  in  Table  VII.  There  are  some  variations  due  to 
individuality  but  in  general  the  percent  decreases  from  birth  to 
four  years.  The  percent  also  decreases  with  increasing  plane  of 
nutrition.  Put  in  different  language  the  percent  of  hide  and  hair 
decreases  with  increasing  age  and  fatness  of  the  animal. 

Blood. — In  Table  VIII  are  presented  the  figures  for  the  per- 
cent of  blood  referred  to  empty  weight.  The  maximum  is  at  three 
months.  At  birth  it  is  less  and  as  age  increases  beyond  three 
months  it  becomes  less.  In  the  poorer  fed  groups  the  percent  of 
blood  becomes  fairly  constant,  however,  while  in  the  full  fed  group 
it  continues  to  decrease  with  age.  In  the  early  stages  the  full  fed 
animals  have  as  much  or  more  than  the  others  but  as  the  age  in- 
creases the  Group  I steers  have  a materially  smaller  percent  of 
blood. 


Table  VII. — Percent  Hide  and  Hair  to 
Empty  Weight. 


Table  VIII. — Percent  Blood  to  Empty 
Weight. 


Age 

At  birth  

3 months  

5%  months  

8 Y2  months  

11  months  

18  months  

21-26  months  

34  months  

38  months  stunted  . . 

40  months  

45  months  

47-48  months  


Group 

Group 

Group 

I 

II 

III 

12.13 

12.13 

12.13 

10.51 

9.48 

9.26 

8.16 

10.60 

9.27 

8.53 

8.62 

9.04 

8.81 

9.72 

9.44 

7.40 

8.69 

8.65 

9.80 

10.47 

7.43 

8.23 

7.15 

5.88 

8.35 

9.34 

5.87 

8.90 

9.61 

6.15 

8.36 

8.81 

Age 

At  birth  

3 months  

5^2  months  

SV2  months  

11  months  

18  months  

21-26  months  

34  months  

38  months  stunted  . . 

40  months  

45  months  

47-48  months  


Group 

Group 

Group 

I 

II 

III 

4.93 

4.93 

4.93 

6.24 

5.35 

6.37 

5.18 

5.25 

5.35 

5.08 

5.85 

5.19 

4.33 

4.54 

5.06 

4.09 

4.93 

4.41 

4.53 

5.13 

4.16 

4.85 

3.61 

3.48 

4.43 

5.28 

3.33 

4.44 

4.67 

3.52 

4.90 

5.22 

For  a somewhat  different  presentation  of  these  and  other  fig- 
ures bearing  on  the  relation  of  blood,  surface,  and  nitrogen  to  the 
animal  the  reader  is  referred  to  two  earlier  publications  from  this 
Station1 

Heart. — The  percent  of  the  heart  referred  to  the  empty  weight 

JP*  F.  Trowbridge,  C.  R.  Moulton,  L.  D.  Haigh,  The  Maintenance  Requirement  of 
Cattle,  Research  Bulletin  18,  Missouri  Agr.  Expt.  Station  (1915). 

C.  R.  Moulton,  Units  of  Reference  for  Basal  Metabolism  and  Their  Interrelations, 
Four.  Biol.  Chem.  XXIV.  299-320  (1916). 


10 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


is  given  in  Table  IX.  The  figures  are  not  notable  but  are  very  small 
and  decrease  with  increasing  age  and  fatness.  There  is,  however, 
very  little  difference  between  Groups  II  and  III. 

Lungs. — The  proportion  of  lungs  and  trachea  to  the  empty 
weight  (Table  X)  increases  during  the  first  few  months  of  life  and 
then  falls  rather  steadily  with  advancing  age.  The  well  fed  group 
has  materially  less  lungs  than  the  other  groups  especially  at  the 
older  ages. 

Central  Nervous  System. — The  percent  of  brain  and  spinal 
cord  is  given  in  Table  XI.  There  is  a rather  steady  and  uniform 
decrease  with  increasing  age  and  fatness.  From  slightly  less  than 
0.9  percent  at  birth  it  decreases  to  0.09  percent  at  maturity  for  the 
fattest  animal. 

Stomachs. — In  contrast  to  the  foregoing  organs  and  parts  the 
stomachs  (Table  XII)  show  an  increasing  proportion  from  birth 
to  8J/2  months  for  Groups  I and  II  and  from  birth  to  two  years  for 


Table  IX. — Percent 

Heart 

TO 

Empty 

Table  X. — Percent  Lungs 

TO 

Empty 

Weight. 

Weight. 

Age 

Group 

Group  Group 

Age 

Group 

Group  Group 

I 

II 

III 

I 

II 

III 

At  birth  

0.55 

0.55 

0.55 

At  birth  . . . 

1.01 

1.01 

1.01 

3 months  

0.53 

0.43 

0.44 

3 months  . . 

1.30 

1.16 

1.32 

5%  months  

0.54 

0.57 

0.50 

5%  months  . 

1.14 

1.10 

1.26 

8%  months  

0.45 

0.46 

0.56 

8%  months  . 

1.09 

1.21 

1.23 

11  months  

0.38 

0.54 

0.49 

11  months  . 

0.87 

1.12 

1.09 

18  months  

0.45 

0.53 

18  months  . 

0.84 

1.00 

21-26  months  

0.34 

0.44 

0.44 

21-26  months 

0.76 

1.00 

0.62 

34  months  

0.40 

0.49 

34  months  . . 

0.54 

0.90 

38  months  stunted  . 

0.32 

38  months  stunted  . 0.67 

40  months  

0.41 

0.39 

0.49 

40  months  . 

0.55 

0.89 

1.07 

45  months  

0.30 

0.44 

0.42 

45  months  . 

0.63 

0.83 

0.84 

47-48  months  

0.27 

0.40 

0.36 

47-48  months 

0.47 

0.79 

0.92 

Table  XI. — Percent 

Brain 

and  Cord  to 

Table  XII.— 

Percent  Stomachs  to  Empty 

Empty  Weight. 

Weight. 

Age 

Group 

Group  Group 

Age 

Group 

Group  Group 

I 

II 

III 

I 

II 

III 

At  birth  

0.86 

0.86 

0.86 

At  birth  . . . 

0.99 

0.99 

0.99 

3 months  

0.41 

0.51 

0.50 

3 months  . . . 

2.01 

1.55 

1.88 

5 x/<i  months  

0.32 

0.47 

0.61 

5%  months 

2.51 

1.95 

2.19 

8%  months  

0.27 

0.37 

0.58 

8%  months 

3.03 

3.17 

2.83 

11  months  

0.20 

0.29 

0.37 

1 1 months  . 

2.73 

2.85 

2.83 

18  months  

0.14 

0.30 

18  months  . 

2.37 

2.78 

21-26  months  

0.15 

0.24 

0.27 

21-26  months 

2.69 

2.77 

3.33 

34  months  

0.11 

0.18 

34  months  . 

1.96 

2.30 

38  months  stunted  . 

0.11 

38  months  stunted  . 1.90 

40  months  

0.09 

0.15 

0.24 

40  months  . 

1.32 

2.57 

2.51 

45  months  

0.10 

0.18 

0.19 

45  months  . 

1.74 

2.44 

2.52 

47-48  months  

0.09 

0.14 

0.20 

47-48  months 

1.74 

2.25 

2.70 

Studies  In  Animal  Nutrition — II 


11 


Group  III.  The  proportion  then  becomes  less  and  continues  to 
fall  for  the  full  fed  group.  In  the  early  months  the  full  fed  group 
has  the  greater  percent  of  stomachs  but  in  later  years  the  smaller 
percent  of  stomachs. 

These  figures  show  a retarded  development  of  the  stomachs 
for  the  group  receiving  the  lightest  ration.  All  groups  show  a 
decrease  in  percent  with  advancing  maturity. 

Intestines. — The  intestines  (Table  XIII),  however,  show  a 
much  less  marked  increase  in  percent  with  the  early  months  and 
a much  less  marked  retarding  of  development  with  Group  III. 
After  the  first  few  months  the  percent  of  intestines  decreases  with 
increasing  age  and  fatness. 

The  above  evidence  may  be  affected  by  the  thickness  or  diam- 


Table  XIII. — Percent  Intestines  to 

Empty  Weight. 


Age  Group  Group  Group 


I 

II 

III 

At  birth  

2.60 

2.60 

2.60 

3 months  

2.75 

2.66 

3.44 

5%  months  

2.37 

2.80 

2.93 

8%  months  

2.61 

2.99 

2.91 

11  months  

1.79 

2.15 

2.37 

18  months  

1.29 

1.92 

21-26  months  

2.00 

1.55 

2.24 

34  months  

1.02 

1.34 

38  months  stunted  . 

0.83 

40  months  

0.64 

0.97 

1.24 

45  months  

0.76 

0.93 

0.99 

47-48  months  

0.60 

1.08 

1.11 

Table  XIV. — Cm.  of  Intestines  per  Ko.  of 
Empty  Weight. 


Age 

Group 

Group  Group 

I 

II 

III 

At  birth  

45.24 

45.24 

45.24 

3 months  

34.83 

37.17 

34.61 

5%  months  

22.03 

31.41 

39.03 

8%  months  

20.78 

27.87 

34.43 

11  months  

15.17 

25.19 

23.83 

18  months  

11.69 

19.21 

21-26  months  

10.88 

13.80 

34  months  

6.99 

9.46 

38  months  stunted  . 

6.05 

40  months  

6.28 

10.37 

11.35 

45  months  

7.18 

10.16 

10.62 

47-48  months  

5.95 

11.23 

10.62 

eter  of  the  intestines.  The  relative  length  is  given  in  Table  XIV. 
It  is  seen  that  at  birth  the  beef  animal  has  much  the  greatest  length 
of  intestines  per  unit  of  empty  weight  and  that  this  proportion  de- 
creases continuously  with  advancing  age.  After  three  months  the 
proportion  decreases  with  increased  plane  of  nutrition  and  conse- 
quent size  and  fatness  of  the  animal. 

At  birth  there  are  45  centimeters  per  kilogram  of  empty  weight 
while  the  oldest  fattest  steer  shows  less  than  six. 

Liver. — The  liver  is  a most  important  chemical  laboratory  of 
the  body.  Its  proportion  should  be  affected  by  the  amount  of  the 
ration  perhaps  more  than  by  the  size  of  the  animal.  The  figures 
given  in  Table  XV  show  a rather  steady  decrease  from  birth  to 
four  years  in  Group  II  and  Group  III.  Group  I,  however,  shows 
an  increase  up  to  8 y2  months  and  then  a decrease  with  increasing 
age. 


12 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


From  birth  up  to  about  two  years  old  the  Group  I cattle  show 
relatively  more  liver  than  do  the  other  groups.  Thereafter  this 
group  shows  less  but  not  more  than  one-third  less  than  the  Group 
III  animals.  The  highest  figure  is  1.84  percent  and  the  lowest  0.73 
percent. 


Table  XV. — Percent  Liver  to  Empty 
Weight. 


Table  XVI. — Percent  Kidneys  to  Empty 
Weight. 


Age 

At  birth  

3 months  

5%  months  

8%  months  

1 1 months  

18  months  

21-26  months  

34  months  

38  months  stunted  . 

40  months  

45  months  

47-48  months  


Group 

Group 

Group 

I 

II 

III 

1.76 

1.76 

1.76 

1.79 

1.49 

1.75 

1.84 

1.19 

1.28 

1.66 

1.65 

1.50 

1.39 

1.25 

1.16 

1.24 

1.15 

1.00 

0.97 

1.08 

0.89 

0.90 

0.84 

0.73 

0.83 

0.94 

0.77 

0.84 

0.99 

0.76 

0.89 

1.14 

Age 

At  birth  

3 months  

5%  months  

8%  months  

11  months  

18  months  

21-26  months  

34  months  

38  months  stunted  . 

40  months  

45  months  

47-48  months  


Group 

Group 

Group 

I 

II 

III 

0.32 

0.32 

0.32 

0.34 

0.68 

0.62 

0.38 

0.32 

0.41 

0.26 

0.31 

0.35 

0.22 

0.31 

0.26 

0.19 

0.26 

0.18 

0.21 

0.25 

0.16 

0.18 

.... 

0.16 

0.16 

0.22 

0.24 

0.13 

0.19 

0.20 

0.13 

0.22 

0.25 

Kidneys. — The  kidneys  (Table  XVI)  show  for  Group  I cat- 
tle much  the  same  changes  as  are  shown  by  the  liver.  There  is  an 
increase  from  birth  to  5^2  months  and  then  a steady  decrease. 
Groups  II  and  III  show  a very  striking  increase  in  proportion  of 
kidneys  at  three  months  where  the  figure  is  double  that  shown  at 
birth.  Thereafter  there  is  a decrease  to  two  years  of  age  when  the 
figures  become  about  constant. 

The  percent  of  kidneys  is  smaller  the  higher  the  plane  of  nu- 
trition and  the  bigger  and  fatter  the  animal.  This  is  especially  true 
after  the  age  of  one  year. 

Spleen. — The  percent  of  spleen  to  empty  weight  is  given  in 
Table  XVII.  It  increases  at  first  with  increasing  age  and  then 
remains  quite  constant  for  Groups  II  and  III.  For  Group  I it  de- 
creases from  three  months  until  38  to  40  months  when  it  becomes 
constant. 

During  the  early  months  the  Group  I cattle  show  as  much,  or 
more,  spleen  as  do  the  other  groups.  Thereafter  the  proportion 
decreases  with  increased  plane  of  nutrition. 

Pancreas. — The  figures  for  the  pancreas  are  presented  in  Table 
XVIII.  In  discussing  these  it  must  be  remembered  that  this  organ 
becomes  embedded  in  fat  as  the  animal  grows  fatter  and  even  ap- 
pears to  have  large  areas  of  fat  scattered  through  it.  The  person 


Studies  In  Animal  Nutrition — II 


13 


making  the  separation  has  increasing  difficulty  in  separating  the 
organ  from  the  fat  or,  in  fact,  in  telling  whether  he  has  obtained 
the  organ  at  all. 


Table  XVII. — Percent  Spleen  to  Empty 
Weight. 


Age 

Group 

I 

Group  Group 
II  III 

At  birth  

0.19 

0.19 

0.19 

3 months  

0.31 

0.26 

0.24 

5%  months  

0.28 

0.29 

0.25 

8*4  months  

0.23 

0.24 

0.22 

11  months  

0.21 

0.21 

0.24 

18  months  

0.19 

0.25 

21-26  months  

0.28 

0.25 

0.21 

34  months  

0.22 

0.27 

38  months  stunted  . 

0.17 

40  months  

0.16 

0.19 

0.24 

45  months  

0.14 

0.21 

0.33 

47-48  months  

0.15 

0.25 

0.26 

Table  XVIII. — Percent  Pancreas  to 
Empty  Weight. 


Age 

Group 

Group  Group 

I 

II 

III 

At  birth  

0.08 

0.08 

0.08 

3 months  

0.10 

0.10 

0.15 

5 *4  months  

0.13 

0.10 

0.10 

8V2  months  

0.12 

0.17 

0.17 

11  months  

0.12 

0.13 

0.13 

18  months  

0.13 

0.16 

21-26  months  

0.06 

0.09 

0.06 

34  months  

0.06 

0.06 

38  months  stunted  . 

0.13 

40  months  

. 0.11 

0.12 

0.14 

45  months  

0.11 

0.13 

0.14 

47-48  months  

0.10 

0.15 

0.15 

With  these  reservations  in  mind  it  may  be  said  that  the  propor- 
tion of  pancreas  increases  with  increasing  age  until  8*4  months  is 
reached.  The  percent  then  becomes  somewhat  less  or  at  least  no 
greater.  There  is  but  little  difference  between  the  groups.  The 
smaller  proportion  shown  after  40  months  may  or  may  not  be  fact 
owing  to  the  difficulties  stated. 

Main  Divisions  of  Animal. — Figure  2 shows  the  growth  of  cer- 
tain tissues  and  parts  of  the  animals  of  all  three  groups  from  birth 
to  four  years.  A general  idea  of  the  weight  and  proportion  of  the 
skeleton,  lean  flesh,  fatty  tissue,  organs,  hide,  and  remainder  can 
be  obtained  from  a study  of  the  figure. 

Figures  3 to  5 show  the  percent  of  the  various  parts  enumer- 
ated above  referred  to  the  empty  animal. 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Fig.  2. — Growth  of  various  divisions  of  the  animal. 


Studies  In  Animal  Nutrition — II 


15 


In  Group  I (Figure  3)  the  skeleton  decreases  from  28  percent 
at  birth  to  10  percent  at  four  years.  The  lean  flesh  is  more  con- 
stant, increasing  from  39  percent  at  birth  to  44.5  percent  at  8^2 
months  while  the  mature  steer  has  but  32  percent.  The  fatty  tissue 


1*1 

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Fig.  3. — Distribution  of  tissues  in  empty  animal — Group  I. 


increases  very  markedly  from  an  amount  so  small  as  to  be  insep- 
arable from  the  lean  at  birth  to  40  percent  at  four  years.  The  or- 
gans of  the  animal  are  about  11  percent  from  birth  to  8 months. 
They  then  decrease  to  about  6 percent  at  four  years.  The  bloorl 
and  the  hide  have  been  discussed  above. 


16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


In  Group  II  (Figure  4)  less  change  with  advancing  age  is 
shown.  This  is  largely  due  to  a smaller  amount  of  fatty  tissue  be- 
ing formed.  The  skeleton  decreases  from  about  28  percent  at  birth 
to  16  percent  at  four  years.  The  lean  flesh  is  about  39  percent  at 
birth  and  increases  to  46  percent  at  45  months.  The  four  year  old 


10 

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Fig.  4. — Distribution  of  tissues  in  empty  animal — Group  II. 


steer  shows  but  41  percent,  probably  an  abnormal  figure  due  to  an 
increase  in  percent  of  fatty  tissue.  The  fatty  tissue  increases  from 
practically  nothing  to  19  percent  at  four  years.  The  organs  are 
about  11  percent  at  birth  and  8.5  percent  at  four  years. 


Studies  In  Animal  Nutrition — II 


17 


Group  III  (Figure  5)  shows  still  less  change.  The  skeleton 
decreases  from  about  28  percent  at  birth  to  18  percent  at  four  years. 
The  lean  flesh  increases  from  39  percent  at  birth  to  45  to  48  percent 
at  the  end.  The  fatty  tissue  is  small  in  amount  running  from  prac- 
tically nothing  at  birth  to  only  a little  over  11  percent  at  four  years. 
The  organs  decrease  from  11  percent  at  birth  to  9 percent  at  four 
years. 


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Fig.  5. — Distribution  of  tissues  in  empty  animal — Group  III. 


Loss  on  Cooling  and  Cutting. — The  weights  and  percents  of 
the  main  divisions  of  the  empty  animal  and  the  calculated  loss  on 
cooling  and  cutting  are  given  in  the  appendix  in  Tables  16  to  20. 
The  losses  vary  from  practically  nothing  to  over  6 percent  in  the 
case  of  one  animal.  Five  animals  show  about  5 percent.  The  other 


18 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


25  animals  show  less  than  3 percent  loss.  This  loss  is  largely 
moisture  lost  from  the  animal  during  slaughter,  cooling,  and  cut- 
ting. While  the  differences  between  the  groups  are  not  large,  in 
general  the  greater  percent  losses  are  from  the  thin  animals.  The 
covering  of  fat  on  the  Group  I steers  protects  the  carcass  and  even 
the  offal  from  as  great  a moisture  loss. 


PISTRIBUTION  or  CUTS  •"  CARCASS  -group*. 

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AGE  IN  MONTHS. 

Fig.  6. — Distribution  of  cuts  in  carsass — Group  I. 


DISTRIBUTION  OF  WHOLESALE  CUTS 

Tables  21  to  25  in  the  appendix  give  the  detailed  figures  for  the 
distribution  of  the  wholesale  cuts  in  the  carcass.  Figures  6 to  8 
show  these  figures  graphically. 

In  Group  I (Figure  6)  the  shank  (hind  shank)  decreases  from 
about  5 percent  at  three  months  to  2 percent  at  four  years.  The 


Studies  In  Animal  Nutrition — II 


19 


round  decreases  from  20  percent  to  13  percent.  The  rump  in- 
creases from  3 to  a little  over  4 percent.  The  flank  increases  from 
3 to  6 percent.  The  kidney  fat  increases  from  about  1 percent  at 
three  months  to  4 percent  in  the  baby  beef  (18  to  21  months)  and 
then  decreases  again  to  2 to  3 percent.  The  loin  increases  from 
16.5  percent  to  21  percent  at  four  years.  The  rib  cut  does  not  vary 


DISTRIBUTION  or  CUTS  »*  CARCASS.  «- CROUP®. 

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Fig.  7. — Distribution  of  cuts  in  carcass — Group  II. 


much  being  8 percent  at  three  months,  and  9 to  10  percent  at  four 
years.  The  plate  increases  from  10  percent  to  17  percent.  The 
chuck  decreases  from  25  percent  to  20  percent.  The  neck  decreases 
from  about  2 percent  to  1 percent  and  the  shin  (fore  shank)  de- 
creases from  6 percent  to  3 percent. 


20 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


The  Group  II  cattle  show  much  less  marked  changes  in  distri- 
bution of  the  cuts  from  three  months  to  four  years.  The  shank 
decreases  from  5 to  3 percent,  the  round  from  20  to  18  percent,  the 
shin  from  6 to  4 percent,  and  the  neck  from  nearly  2 to  1 percent. 
The  flank  increases  from  2 to  4 percent,  the  loin  increases  from  16 
to  18  percent  at  Sy2  months  and  then  decreases  again  to  less  than 
17  percent,  and  the  plate  increases  from  less  than  9 percent  to 
nearly  14  percent.  The  rump,  kidney  and  kidney  fat,  rib,  and 
chuck  remain  fairly  constant. 

In  Group  III  (Figure  8)  there  is  less  change  with  age.  The 
shank,  kidney  and  kidney  fat,  and  shin  decrease  with  age.  The 
loin  and  plate  show  fair  increases  while  the  rump  and  flank  show 
slight  increases.  The  round,  rib,  chuck,  and  neck  show  individual 
variations  but  on  the  whole  are  constant. 

The  fatter  animal,  then,  increases  its  proportion  of  loin,  the 
most  expensive  cut,  of  rump,  a less  expensive  cut,  and  of  flank 
and  plate  very  cheap  cuts.  The  rib  cut,  a rather  expensive  cut,  in- 
creases but  slightly.  On  the  average  for  all  three  groups  it  is  con- 
stant. The  round,  a valuable  cut,  decreases  with  increasing  fatness 
as  do  the  chuck  and  the  neck.  The  shin  and  shank  decrease  in  all 
cases  with  increasing  age  irrespective  of  fattening.  Summing  up 
the  changes  with  respect  to  the  effect  of  fattening  on  the  three  ex- 
pensive cuts — loin,  rib,  and  round — it  is  seen  that  the  first  increases 
with  fattening,  the  second  remains  fairly  constant,  and  the  third 
decreases. 

LEAN,  FAT,  AND  BONE  IN  THE  CARCASS 

Figure  9 (Tables  26  to  30  in  the  appendix)  shows  the  propor- 
tions of  skeleton,  lean  flesh  and  fatty  tissue  in  the  entire  carcasses 
of  the  steers  from  three  months  to  four  years  for  all  three  groups. 

In  the  Group  I steers  the  skeleton  decreases  from  about  25 
percent  to  10  percent,  and  the  lean  flesh  decreases  from  67  percent 
to  42  percent.  The  fatty  tissue  of  the  carcass,  on  the  other  hand, 
increases  from  6 percent  to  nearly  48  percent. 

In  Group  II  the  changes  are  not  so  marked  since  relatively  less 
fattening  occurs.  The  skeleton  decreases  from  nearly  28  percent 
to  a little  over  18  percent  and  the  lean  flesh  decreases  from  66  per- 
cent at  three  months  and  68  percent  at  11  and  26  months  to  58  per- 
cent at  four  years.  The  fatty  tissue  increases  from  5 percent  to  22 
percent. 


Studies  In  Animal  Nutrition — II 


21 


The  changes  are  still  less  marked  in  Group  III.  The  skeleton 
decreases  from  27  percent  to  a little  over  20  percent  while  the  lean 
flesh  increases  from  67  percent  at  three  months  to  about  70  percent 


5_| 

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Fig.  8. — Distribution  of  cuts  in  carcass — Group  III. 


from  18*/2  months  on  to  45  months.  The  four-year-old  steer  shows 
a decrease  to  a little  over  66  percent.  The  fatty  tissue  increases 
from  a little  over  3 percent  to  over  12  percent. 


22 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


PISTRiBUTIOM  of  lea « . FAT,  ais*  f ONE  •«  CAF 

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Fig.  9. — Distribution  of  lean,  fat,  and  bone  in  carcass. 


Studies  In  Animal  Nutrition — II 


23 


DISTRIBUTION  OF  THE  TOTAL  LEAN  FLESH 

Figures  10  to  12  (Tables  31  to  35  in  the  Appendix)  show  the 
distribution  of  the  total  lean  flesh  of  the  animal  among  the  whole- 
sale cuts. 


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AC t IN  MONTHS. 

Fig.  10. — Distribution  of  lean  in  cuts  of  carcass — Group  I. 


In  Group  I the  head  and  tail  contain  about  2 to  3 percent  of 
the  total  lean.  The  shin  and  shank  together  have  5 to  6 percent. 
The  chuck  and  neck  combined  have  26  to  29  percent.  The  plate 
has  10  to  14  percent,  the  rib  8 to  10  percent,  the  loin  17  to  19  per- 
cent, the  flank  3 to  4 percent,  the  rump  2 to  3 percent,  and  the 
round  23  to  19.5  percent.  The  shin  and  shank,  the  head  and  tail, 


24 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


and  the  round  contain  a smaller  part  of  the  total  lean  with  advanc- 
ing age  and  fatness  of  this  group.  The  chuck  and  neck,  rib,  loin, 
flank,  and  rump  show  but  little  effect  of  age  and  fatness.  The  plate, 
on  the  other  hand,  shows  an  appreciable  relative  increase  with  ad- 
vancing age  and  fatness. 


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AGE.  IN  MONTHS. 

Fig.  11. — Distribution  of  lean  in  cuts  of  carcass — Group  II. 


In  Group  II  (Figure  11)  the  head  and  tail,  shin  and  shank, 
rib,  flank,  and  rump  have  about  the  same  percent  of  the  total  lean 
as  shown  by  Group  I.  The  chuck  and  neck  and  the  round  contain 
relatively  more  in  the  Group  II  steers,  while  the  plate  and  loin  con- 
tain relatively  less.  The  differences  are,  however,  not  very  great 
in  any  case  being  generally  within  2 percent.  The  plate  shows 


Studies  In  Animal  Nutrition — II 


25 


an  increase  with  increasing  age  while  the  round  shows  a decrease. 
The  other  cuts  show  little  effect  of  age. 

With  the  Group  III  animals  (Figure  12)  the  head  and  tail, 
shin  and  shank,  flank,  and  rump  have  about  the  same  proportion 
of  total  lean  as  shown  by  the  other  groups.  The  head  and  tail  in 


20 

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AGC  IN  MONTHS. 

Fig.  12. — Distribution  of  lean  in  cuts  of  carcass — Group  III. 


some  cases  runs  1 percent  higher  and  the  shin  and  shank  2 percent 
higher.  The  plate,  rib,  loin,  and  round  vary  over  wider  limits  while 
the  chuck  and  neck  keep  within  narrower  limits.  The  round  varies 
from  20  to  26  percent,  the  loin  from  14.5  to  19  percent,  the  plate 
from  7 to  12  percent,  and  the  rib  from  7 to  10  percent.  There  seems 
to  be  no  consistent  effect  of  age  excepting  a reduction  of  the  head 
and  tail  and  the  shin  and  shank. 


26 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


DISTRIBUTION  OF  TOTAL  FAT  FLESH 

The  distribution  of  the  total  fatty  tissue  exclusive  of  the  offal 
fat  is  shown  in  Figures  13  to  15  (Tables  36  to  40  in  the  Appendix). 
There  is  much  greater  variation,  and  age  and  fatness  affect  the 
distribution  of  fatty  tissue  quite  markedly. 

In  Group  I (Figure  13)  the  head  and  tail  contains  3 to  4 per- 
cent of  the  fatty  tissue  in  the  young  animal  and  less  than  0.5  per- 
cent in  the  four-year-old  steer.  The  shin  and  shank  start  with 
about  6 percent  and  have  less  than  2 percent  at  four  years.  The 
chuck  and  neck  have  about  19  percent  at  three  months  and  13  to 
15  percent  at  four  years.  The  plate  increases  from  7.5  percent  to 
21  percent,  and  the  rib  increases  from  2 to  10  percent.  The  kidney 
fat  increases  from  10  percent  at  three  months  to  16  percent  in  the 
baby  beef  (11  to  18  months)  and  drops  again  to  6 or  7 percent  in 
the  old  animals.  The  loin  increases  its  proportion  from  20  percent 
to  25  percent.  The  flank  decreases  from  11  to  9 percent  and  the 
round  decreases  markedly  from  17  to  about  7 percent.  The  rump 
increases  slightly  from  3.5  to  5 or  6 percent.  With  increasing  age 
and  fatness,  therefore,  a greater  part  of  the  fatty  tissue  is  found 
in  the  plate,  rib,  loin,  and  rump  and  a smaller  part  in  the  head  and 
tail,  shin  and  shank,  chuck  and  neck,  flank,  and  round.  The  kidney 
fat  at  first  increases  and  then  decreases. 

In  Group  II  (Figure  14)  the  head  and  tail  have  5 to  6 percent 
of  the  total  fat  in  the  young  animal  and  only  1 percent  in  the  old 
animal.  The  shin  and  shank  decrease  from  8 to  2 percent.  The 
chuck  and  neck  starts  at  about  18  percent.  It  then  rises  to  21  per- 
cent at  11  months  and  falls  to  about  13  percent  at  34  months.  It 
then  rises  again  to  21  percent  at  44^4  months  and  decreases  to  15 
percent  at  four  years.  The  plate  increases  its  proportion  of  fat 
from  9 to  20  percent  with  advancing  age.  The  rib  has  none  of  the 
total  fat  at  three  months  and  13  percent  at  four  years.  The  kidney 
fat  is  9 to  10  percent  of  the  total  from  three  months  to  34  months 
with  the  exception  of  the  11-month-old  steer  which  has  but  5.5 
percent.  From  40  months  on  there  is  but  6 percent  of  the  total 
here.  The  loin,  like  the  chuck  and  neck,  shows  two  maxima.  It 
increases  from  16  percent  at  three  months  to  26  percent  at  8^4 
months.  It  then  decreases  to  21  percent.  At  34  to  40  months  it  is 
23  percent,  only  to  decrease  to  less  than  21  percent  at  four  years. 
The  flank  in  general  shows  a decrease  in  the  proportion  of  total 
fat  found  here.  The  variations  in  the  rump  are  not  a function  of 


PERCENT  or  TOTAL  FAT  FLESH. 


Studies  In  Animal  Nutrition — II 


FMSTRI&UTION  Of  FAT  FLESH 


GROUP  I, 


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TAIL. 


MONTHS. 


Fig.  13. — Distribution  of  fat  in  cuts  of  carcass — Group  1. 


28 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


age,  while  the  round  decreases  from  19  percent  to  10  percent  at  40 
months  and  then  increases  again  to  13  percent.  With  increasing 
fatness,  then,  a greater  part  of  the  total  fatty  tissue  is  found  in 
the  plate  and  rib.  This  is  true  in  general  of  the  loin,  although 


PISTRIBUTIOM  or  FAT  FLESH  :•  groupie. 

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ACE  IN  MONTHS 

Fig.  14. — Distribution  of  fat  in  cuts  of  carcass — Group  II. 


there  are  cycles  which  have  higher  maxima  at  younger  ages.  The 
chuck  and  neck  show  cycles  also  but  have  less  at  the  end.  The 
rump  and  flank  show  little  change  with  age,  while  the  head  and 
tail,  shin  and  shank,  and  round  decrease. 


Studies  In  Animal  Nutrition — II 


29 


Group  III  (Figure  15)  shows  greater  individual  variations  than 
do  the  other  groups.  Practically  every  wholesale  cut  shows  irregu- 


2o 

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AGE ’IN  MONTHS. 

it  Al  48. 

Fig.  15. — Distribution  of  fat  in  cuts  of  carcass — Group  III. 


lar  changes.  The  head  and  tail  and  the  shin  and  shank  decrease 
rather  uniformly  while  the  plate  and  rib  increase  uniformly.  The 
round  has  less  at  four  years  than  at  three  months  while  the  loin 


30 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


has  more,  but  there  are  many  ups  and  downs  in  between.  The 
changes  in  the  other  cuts  do  not  consistently  follow  age. 

DISTRIBUTION  OF  TOTAL  SKELETON 

Figures  16  to  18  (Tables  41  to  45  in  the  Appendix)  show  the 
changes  in  the  distribution  of  the  total  skeleton  for  the  three  groups 


PISTRISUTIOM  of  SKELETON  •*  croup x. 

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AGE  IN  MONTHS. 

Fig.  16. — Distribution  of  bone  in  cuts  of  carcass — Group  1. 


with  advancing  age.  The  values  are  much  more  constant  than  for 
either  the  lean  flesh  or  the  fatty  tissue. 

In  Group  I (Figure  16)  the  head  and  tail  have  about  13  percent 


Studies  In  Animal  Nutrition — II 


31 


of  the  total  skeleton  with  variations  from  about  11.5  to  14.5.  The 
feet  bones  increase  from  about  12.5  percent  at  three  months  to  15 
percent  at  8 months  and  then  decrease  to  9.5  percent  at  four 
years.  The  shin  and  shank  decrease  rather  uniformly  from  17.5 


PISTRJSUTiON  ©?  SKELETON  *•  oroupst. 

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AGE  IN  M0NTN5. 

Fig.  17. — Distribution  of  bone  in  cuts  of  carcass — Group  II. 


to  15.5  percent.  The  chuck  and  neck  bones  increase  from  18  to  21 
percent  at  18  months  and  then  decrease  to  19.5  percent.  The  plate 
bones  vary  from  7.5  to  9 percent  increasing  slightly  with  age.  The 
rib  varies  around  8 percent  and  the  loin  around  11  percent.  The 
flank  has  but  a tip  of  rib  bone  in  it  and  varies  from  a few  hun- 


32 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


dredths  of  a percent  to  about  0.2  percent  increasing  with  age.  The 
rump  increases  slightly  from  3 to  5 percent,  while  the  round  de- 
creases very  slightly  from  9.5  to  9 percent. 


PI5TRIBUTIC 

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or  SKELETON  » growpES. 

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AGE.  IN  MONTHS. 

48 

Fig.  18. — Distribution  of  bone  in  cuts  of  carcass — Group  III. 


Groups  II  and  III  (Figures  17  and  18)  show  somewhat  greater 
variations  but  the  range  of  the  figures  is  much  the  same.  The  in- 
crease in  the  proportion  of  skeleton  found  in  the  feet  at  8^2  months 
is  not  seen.  In  other  respects  the  figures  are  much  as  those  for 
Group  I. 


Studies  In  Animal  Nutrition — II 


33 


DISTRIBUTION  OF  LEAN,  FAT,  AND  BONE  IN  THE 
WHOLESALE  CUTS 

The  Shin  (Fore  Shank). — With  the  Group  I animals  (Figure 
19)  the  lean  forms  about  50  percent  of  the  shin,  the  fat  5 to  18  per- 
cent and  the  bone  48  to  34  percent.  The  percent  of  fat  increases 
with  age  and  fatness  and  the  bone  decreases,  while  the  lean  varies. 

With  the  Group  II  animals  the  lean  forms  44  to  53  percent  of 
the  cut,  the  fat  3 to  6 percent,  and  the  bone  50  to  40  percent.  The 
lean  increases  with  age  while  the  bone  decreases.  The  fat  changes 
but  little  with  age. 

With  the  thinnest  animals — Group  III — the  lean  is  46  to  57 
percent  of  the  cut,  the  fat  3 to  5 percent,  and  the  bone  50  to  40  per- 
cent. There  are  more  irregularities  shown  in  this  group,  but  in 
general  the  lean  increases  in  percent  with  age  while  the  bone  de- 
creases. 

The  Neck. — The  neck  of  the  Group  I animals  (Figure  20)  con- 
tains from  66  to  40  percent  lean,  12  to  33  percent  fat,  and  18  to  30 
percent  bone.  The  fat  increases  with  increasing  age  and  fatness 
while  the  lean  decreases.  The  bone  varies  between  the  limits  given 
without  respect  to  age.  The  amount  and  composition  of  this  cut 
will  depend  much  upon  the  general  conformation  and  fatness  of 
the  carcass.  There  will  then  be  considerable  differences  between 
individuals. 

In  the  Group  II  animals  the  neck  is  50  to  67  percent  lean,  4 to 
14  percent  fat,  and  23  to  39  percent  bone.  The  percent  of  lean,  of 
fat,  or  of  bone,  does  not  seem  to  depend  on  age. 

The  Group  III  animals  show  even  greater  variations  and  no 
better  correlation  between  age  and  composition. 

The  Chuck. — The  chuck  (Figure  21)  of  the  Group  I animals 
has  72  to  58  percent  lean,  4 to  30  percent  fat  and  23  to  12  percent 
bone.  The  lean  and  bone  decrease  with  age  while  the  fat  increases. 

With  the  Group  II  animals  the  lean  is  70  to  75  percent  of  the 
cut,  the  fat  3 to  12  percent  and  the  bone  26  to  17  percent.  The  fat 
increases  with  age,  the  bone  decreases,  and  the  lean  is  about  con- 
stant. 

In  the  case  of  the  Group  III  animals  the  lean  runs  70  to  75 
percent,  the  fat  2 to  8 percent  and  the  bone  24  to  19  percent.  The 
fat  increases  slightly  with  age,  the  bone  decreases  slightly  while 
the  lean  is  fairly  constant. 

The  Plate. — The  plate  (Figure  22)  becomes  rather  a fat  cut 


34 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Fig.  19. — Distribution  of  lean,  fat  and  bone  in  the  shin. 


PERCENT  of  WHOLESALE  CUT. 


Studies  In  Animal  Nutrition — II 


35 


LEAN  FAT  AMD  BONE  IN  THE  NECK. 


10 

” 

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BONE. 

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FAT. 


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60 

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r-j 

10 

0 

LEAN. 

3 SK  8’*  U A&£  IQ  21  1(1  MOMTMS-  35  33\  44S  47 


Fig.  20. — Distribution  of  lean,  fat  and  bone  in  the  neck. 


36 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


10 

L£ 

IA 

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F 

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0-10 

BONE. 

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£ io 

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60 

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3 5^  & I1  AGE  18  21  IN  M0NTHS.3S  39*  44*47 

Fig.  21. — Distribution  of  lean,  fat  and  bone  in  the  chuck. 


Studies  In  Animal  Nutrition — II 


37 


l 

.E 

1A? 

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NF 

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60 

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i 44S 

.49 

Fig.  22. — Distribution  of  lean,  fat  and  bone  in  the  plate. 


38 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


with  the  Group  I animals.  The  lean  is  6.9  to  35  percent  of  the  cut, 
the  fat  5 to  58  percent,  and  the  bone  25  to  7 percent. 

The  Group  II  animals  show  less  extensive  changes.  The  lean 
runs  65  to  60  percent  except  in  the  case  of  the  oldest  animal  where, 
on  account  of  a large  increase  in  fat,  it  drops  to  50  percent.  The 
fat  runs  from  about  6 to  35  percent  and  the  bone  28  to  17  percent. 
The  lean  decreases  slightly  with  age.  The  bone  decreases  with 
age  while  the  fat  increases. 

With  the  Group  III  steers  the  lean  runs  69  to  63  percent,  the 
fat  2 to  19  percent,  and  the  bone  29  to  17  percent.  There  is  a very 
slight  decrease  in  percent  lean  with  increasing  age.  The  fat  in- 
creases while  the  bone  decreases  with  increasing  age. 

The  Rib. — Figure  23  shows  the  composition  of  the  rib.  With 
the  Group  I steers  the  lean  runs  from  66  to  37  percent,  the  fat  from 
2 to  51  percent,  and  the  bone  from  33  to  11  percent.  The  lean  and 
bone  decrease  while  the  fat  greatly  increases  with  increasing  age. 

With  the  Group  II  animals  the  changes  are  less  extreme.  The 
lean  runs  from  68  to  58  percent,  the  fat  from  nothing  to  19  percent, 
and  the  bone  from  about  32  to  22  percent.  The  lean  and  bone  de- 
crease with  increasing  age  while  the  fat  increases. 

The  Group  III  animals  show  a fairly  constant  percent  of  lean, 
65  to  70  percent.  The  fat  runs  from  nothing  to  9 percent  and  the 
bone  from  33  to  21  and  25  percent.  The  fat  increases  slightly  with 
age  while  the  bone  decreases. 

The  Loin. — The  loin  is  another  cut  that  becomes  very  fat  with 
increasing  age.  The  Group  I steers  (Figure  24)  have  from  69  to 
37  percent  lean,  8 to  57  percent  fat  and  23  to  7 percent  bone.  The 
lean  and  bone  decrease  while  the  fat  increases  with  age. 

With  the  Group  II  animals  the  lean  decreases  from  70  percent 
to  57  and  the  bone  from  23  to  about  14  percent.  The  fat  on  the 
other  hand  increases  from  5 to  25  percent. 

The  Group  III  steers  show  similar  changes.  The  lean  de- 
creases from  74  to  67  percent  and  the  fat  increases  from  4 to  18 
percent.  The  bone  increases  from  20  to  25  percent  and  then  drops 
to  15  to  17  percent.  The  steer  40 months  old  seems  to  be  abnor- 
mal in  composition. 

The  Flank. — The  flank  becomes  the  fattest  cut  of  them  all 
(Figure  25).  The  Group  I animals  have  in  this  cut  from  69  to 
24  percent  lean  and  from  27  to  75  percent  fat.  The  amount  of  bone 
is  insignificant  being  generally  below  one  percent.  With  the  Group 


Studies  In  Animal  Nutrition — II 


39 


LE 

:a 

M FAT  AMD  BOME  IM  THE  RIB. 

1° 

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3 5%  854  U AGE  18 

Z1  IN  MONTHS 35  39%  44% 47 

Fig.  23. — Distribution  of  lean,  fat  and  bone  in  the  rib. 


40 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


LEA? 

20 

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21  IN  MONTHS. 35  39*1  44k4-7 

Fig.  24. — Distribution  of  lean,  fat  and  bone  in  the  loin. 


Studies  In  Animal  Nutrition — II 


41 


Fig.  25. — Distribution  of  lean,  fat  and  bone  in  the  flank. 


42 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


II  steers  the  lean  decreases  from  72  percent  to  33  percent  while 
the  fat  increases  from  25  to  65  percent.  With  the  Group  III  steers 
the  lean  decreases  from  about  83  to  60  percent,  while  the  fat  in- 
creases from  14  to  38  percent.  In  all  groups  then  the  fat  increases 
with  age  while  the  lean  decreases. 

The  Rump  . — The  changes  in  the  composition  of  the  rump  are 
shown  in  Figure  26.  With  the  Group  I animals  the  lean  decreases 
from  59  to  about  30  percent  while  the  fat  increases  from  8 to  56 
percent.  The  bone  decreases  from  33  to  14  percent. 

With  the  Group  II  animals  with  one  striking  exception  the 
lean  decreases  with  age  running  from  50  to  56  percent  down  to 
44  percent.  The  fat  increases  from  8 to  32  percent,  while  the  bone 
decreases  from  41  to  23  percent. 

In  the  case  of  the  Group  III  steers  the  lean  decreases  from 
about  58  percent  to  49  percent,  and  the  fat  increases  from  5 to  21 
percent.  The  bone  increases  at  first  from  36  to  39  percent  and  then 
decreases  to  28  percent. 

The  Round. — The  round  shows  rather  small  relative  changes 
(Figure  27).  The  Group  I steers  have  from  78  to  63  percent  lean, 
from  5 to  28  percent  fat,  and  from  16  to  9 percent  bone.  The  Group 

11  animals  have  about  80  percent  lean  in  all  cases  but  the  oldest. 
The  fat  runs  from  5 to  16  percent  and  the  bone  from  17  to  about 

12  percent.  In  Group  III  the  lean  runs  between  77  and  81  percent, 
the  fat  increases  from  4 to  9 percent  and  the  bone  decreases  from 
18  to  about  12  percent. 

The  Shank. — The  shank  also  shows  small  relative  changes  in 
make  up.  Figure  28  gives  the  distribution.  The  Group  I steers  have 
about  30  percent  lean  with  a few  animals  running  higher.  The  fat 
increases  from  2 to  16  percent,  but  the  next  to  the  oldest  shows 
22.5  percent.  The  bone  decreases  from  66  to  about  50  percent. 

With  the  Group  II  steers  the  lean  is  fairly  constant  after  11 
months  at  about  36  percent.  At  3 months  it  is  29  percent.  The  fat 
is  small  and  increases  irregularly  with  age.  The  bone  decreases 
from  69  percent  to  a value  running  around  60  percent  from  11 
months  on. 

The  Group  III  animals  show  much  the  same  thing  as  the  Group 
II  steers. 

This  is  a very  bony  cut  and  after  8^4  months  it  is  not  much  af- 
fected by  age. 


PERCENT  OF  WHOLESALE  CUT. 


Studies  In  Animal  Nutrition — II 


43 


LEAN  FAT  AND  BONE  IN  THE  RUMP. 


— 

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BONE. 

20 


to 

n 

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Fig.  26. — Distribution  of  lean,  fat  and  bone  in  the  rump. 


44 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


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Fig.  27. — Distribution  of  lean,  fat  and  bone  in  the  round. 


PERCENT  of  WHOLESALE  CUT. 


Studies  In  Animal  Nutrition — II 


45 


LEAN  FAT  ANP  BONE.  IN  THE  SHANK. 


60 

— 

Aq 

GROUP 

m. 

2o 

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5% 

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1 MONTHS. 

48 

40 

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xc. 

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o. — ■!  1 1 ii — i fat. r — i rn r — -i □ i i 

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AGE  IN  MONTHS. 

AO 

pi 

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X. 

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10 

0 i — 1 1 1 

n rn 

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3o 

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ft  m i 

a- 

AGE.  IN  MONTHS. 


Fig.  28. — Distribution  of  lean,  fat  and  bone  in  the  shank. 


46 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


SUMMARY 

Hereford-Shorthorn  beef  steers  were  fed  on  three  planes  of 
nutrition : Group  I,  full  fed  from  birth ; Group  II,  fed  for  maximum 
growth  without  fattening  giving  gains  of  one  pound  per  day  for 
the  first  two  years ; Group  III,  fed  for  scanty  or  retarded  growth 
gaining  about  .69  pounds  per  day  for  the  first  two  years.  The 
slaughter  house  data  for  31  animals  are  presented. 

With  these  animals  the  ratio  of  empty  weight  to  live  weight 
is  greatest  at  birth,  least  at  8 y2  months,  and  intermediate  at  4 
years.  A lower  plane  of  nutrition  gives  a lower  ratio. 

The  ratio  of  carcass  to  live  weight  decreases  to  8 y2  months  then 
increases  to  a maximum  at  3 to  4 years.  The  better  fed  animals 
have  the  greater  percent.  On  the  empty  weight  basis  the  carcass 
continuously  increases  to  3 to  4 years. 

The  proportions  of  hide  and  hair,  heart,  brain  and  spinal  cord, 
and  intestinal  length  to  empty  weight  decrease  with  increasing 
age  and  fatness.  The  blood,  lungs,  stomachs,  intestines  (weight), 
liver,  kidneys,  spleen,  and  pancreas  increase  relatively  during  the 
early  months  and  then  decrease.  The  maxima  occur  generally 
at  8 y2  months.  The  stomachs  and  liver  show  marked  retardation 
on  the  low  planes  of  nutrition. 

The  proportions  of  skeleton  and  of  total  organs  are  greatest 
at  birth  and  the  total  fleshy  parts  at  4 years. 

In  the  carcass  the  proportions  of  loin,  rump,  flank,  and  plate 
increase  with  increasing  age  and  fatness  of  the  steers.  The  rib 
changes  but  little  while  the  round,  chuck  and  neck,  and  shin  and 
shank  decrease. 

The  distribution  of  the  total  lean  flesh  is  but  little  affected  by 
age  and  fatness  excepting  that  there  is  a slight  reduction  in  the 
proportion  found  in  the  shin,  shank,  head  and  tail  and  in  some  cases 
in  the  round.  The  plate  shows  a larger  part  of  the  total  as  the 
animal  grows  and  fattens. 

The  proportion  of  the  total  fat  flesh  found  in  the  plate,  rib, 
loin,  and  in  some  cases  in  the  rump  increases  with  increasing  fat- 
ness. 

Age  and  fatness  influence  the  distribution  of  the  total  skele- 
ton but  slightly.  The  feet  bones  tend  to  increase  relatively  up  to 
8 Yz  months  (Group  I)  and  then  decrease.  The  chuck  and  neck 


Studies  In  Animal  Nutrition — II 


47 


bones  show  a similar  phenomenon  with  the  maximum  at  18  months. 
The  rump  shows  a slight  increase. 

The  composition  of  the  wholesale  cuts  of  meat  is  affected  by 
increasing  age  and  fatness.  In  general  the  percent  of  fatty  tissue 
increases  and  the  percent  of  bone  decreases.  The  percent  of  lean 
flesh  may  increase,  remain  constant,  or  decrease,  but  on  the  aver- 
age it  decreases.  The  increases  in  percent  of  fat  are  greatest  in  the 
plate,  loin,  and  flank.  The  rib  and  rump  show  rather  large  in- 
creases. 


48 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


APPENDIX 


Table  1. — Slaughter  House  Weights  of  Offal  Parts  (.in  Grams). 


Steer 

556 

554 

555 

557 

552 

548 

3 

3 

3 

5 Months 

5 Months 

5 Months 

Age 

Months 

Months 

Months 

17  Days 

7 Days 

9 Days 

Group 

1 

2 

3 

1 

2 

3 

Live  weight 

111,489 

87,456 

84,725 

204,934 

116,491 

99,255 

Blood 

6,124 

4,197 

4,529 

8,952 

5,219 

4,603 

Heart,  pericardium,  arteries 

993 

591 

554 

2,342 

1,056 

753 

Heart,  marketable 

521 

335 

315 

928 

561 

427 

Heart,  lean 

432 

319 

295 

760 

561 

385 

Lungs  and  trachea 

1,272 

907 

937 

1,972 

1,096 

1,084 

Brain 

275 

273 

252 

351 

332 

374 

Spinal  cord 

123 

122 

101 

200 

134 

152 

Tongue,  including  bones  and  larynx 

884 

660 

432 

908 

673 

571 

Tongue,  marketable 

466 

358 

247 

650 

464 

428 

Tongue  bones,  including  larynx 

106 

142 

53 

131 

96 

88 

Gullet 

209 

146 

193 

189 

355 

204 

Stomachs 

1,975 

1,208 

1,338 

4,341 

1,938 

1,885 

Rumen 

999 

616 

745 

2,614 

1,100 

975 

Reticulum 

260 

147 

147 

358 

188 

203 

Omasum 

391 

161 

176 

724 

374 

363 

Abomasum 

325 

284 

270 

645 

276 

344 

Intestines,  small 

1,944 

1,474 

1,651 

2,863 

1,781 

1,660 

Intestines,  large 

750 

603 

792 

1,233 

1,003 

861 

Intestines, small;  length,  cm 

2,878 

2,452 

2,080 

3,139 

2,630 

2,806 

Intestines,  large;  length,  cm 

540 

450 

380 

667 

491 

550 

Neck  sweetbread 

173 

193 

129 

359 

143 

109 

Heart  sweetbread 

155 

143 

30 

264 

122 

59 

Spleen 

300 

202 

167 

485 

284 

215 

Pancreas 

96 

76 

106 

225 

101 

85 

Liver 

1,760 

1,166 

1,240 

3,003 

1,181 

1,104 

Gall  bladder  and  gall 

25 

23 

8 

148 

55 

34 

Gall 

87 

40 

Kidneys 

338 

530 

439 

649 

316 

353 

Urinary  bladder 

46 

58 

63 

76 

41 

53 

Penis 

110 

116 

59 

155 

94 

79 

Diaphragm 

166 

70 

51 

199 

92 

98 

Caul  fat  

417 

170 

107 

1,981 

363 

Stomach  fat 

146 

73 

20 

1,486 

469 

275 

Intestinal  fat 

839 

401 

335 

3,290 

952 

570 

Hide  and  hair 

10,314 

7,400 

6,580 

14,100 

10,532 

8,328 

Dewclaws 

32 

28 

24 

64 

66 

36 

Teeth 

218 

225 

190 

261 

278 

264 

Horns  

9 

46 

36 

40 

Hoofs 

358 

301 

274 

585 

448 

359 

Right  fore  foot  and  hoof 

761 

666 

610 

1,035 

828 

728 

Left  fore  foot  and  hoof 

760 

653 

630 

1,007 

807 

728 

Right  hind  foot  and  hoof 

719 

651 

596 

1,005 

882 

740 

Left  hind  foot  and  hoof 

707 

734 

595 

1,096 

792 

659 

Fore  quarter,  right 

15,436 

13,096 

12,075 

28,483 

16,023 

13,944 

Bind  quarter,  right 

14,849 

12,358 

10,596 

27,438 

15,135 

12,764 

Left  half 

30,129 

24,675 

22,764 

54,150 

30,580 

27,275 

Studies  In  Animal  Nutrition — II 


49 


Table  2. — Slaughter  House  Weights  of  Offal  Parts  (in  Grams). 


Steer ._,>r  • 

547 

550 

558 

541 

538 

540 

Age 

8 Months 
5 Days 

8 Months 
14  Days 

8 Months 
12  Days 

10  Months 
22  Days 

10  Months 
26  Days 

11  Months 
2 Days 

Group 

1 

2 

3 

1 

2 

3 

Live  weight 

206,175 

147,202 

108,191 

323,836 

180,930 

158,131 

Blood 

8,711 

7,080 

4,666 

12,470 

7,219 

6,967 

Heart,  pericardium,  arteries 

1,624 

1,163 

971 

2,646 

1,670 

1,436 

Heart,  marketable 

771 

556 

506 

1,087 

873 

673 

Heart,  lean 

666 

481 

425 

940 

698 

572 

Lungs  and  trachea 

1,868 

1,460 

1,111 

2,387 

1,825 

1,501 

Brain 

346 

319 

373 

383 

343 

361 

Spinal  cord 

113 

124 

147 

185 

144 

151 

Tongue,  including  bones  and  larynx 

800 

834 

650 

2,336 

1,513 

1,547 

Tongue,  marketable 

586 

523 

464 

1,209 

779 

811 

Tongue  bones,  including  larynx 

161 

139 

111 

309 

221 

170 

Gullet 

384 

209 

331 

538 

400 

324 

Stomachs 

5,190 

3,833 

2,550 

7,876 

4,521 

3.894 

Rumen 

3,127 

1,914 

1,329 

4,050 

2,209 

1,966 

Reticulum 

373 

366 

253 

866 

751 

470 

Omasum 

1,072 

1,015 

579 

1,885 

1,013 

878 

Abomasum 

618 

538 

389 

1,075 

548 

480 

Intestines,  small 

2,649 

2.303 

1,599 

3,922 

2,327 

1,954 

Intestines,  large 

1,826 

1,321 

1,018 

1,322 

1,319 

1,305 

Intestines,  small;  length,  cm 

2,900 

2,755 

2,568 

3,343 

3,393 

2,510 

Intestines,  large;  length,  cm 

663 

620 

531 

716 

610 

772 

Neck  sweetbread 

372 

189 

73 

396 

252 

194 

Heart  sweetbread 

319 

147 

59 

344 

229 

214 

Spleen 

391 

291 

193 

596 

331 

331 

Pancreas 

200 

200 

153 

390 

208 

180 

Liver 

2,851 

1,992 

1,352 

3,832 

1,978 

1,593 

Gall  bladder  and  gall 

138 

117 

28 

275 

153 

83 

Gall 

103 

86 

7 

202 

122 

58 

Kidneys 

450 

379 

318 

645 

487 

363 

Urinary  bladder 

83 

60 

48 

153 

123 

134 

Penis 

181 

112 

149 

226 

146 

131 

Diaphragm 

198 

170 

116 

176 

131 

136 

Caul  fat 

1,118 

532 

150 

4,212 

939 

578 

Stomach  fat 

851 

781 

248 

1,803 

799 

469 

Intestinal  fat 

1,910 

1,297 

431 

4,322 

1,714 

1,260 

Hide  and  hair 

14,618 

10,440 

8,138 

26,576 

15,342 

12,994 

Dewclaws 

86 

53 

38 

99 

65 

49 

Teeth 

310 

228 

274 

304 

240 

278 

Horns 

77 

112 

31 

468 

250 

304 

Hoofs 

574 

384 

374 

770 

570 

432 

Right  fore  foot  and  hoof 

1,177 

820 

764 

1,270 

931 

803 

Left  fore  foot  and  hoof 

999 

780 

719 

1,303 

935 

813 

Right  hind  foot  and  hoof 

1,185 

778 

771 

1,368 

937 

805 

Left  hind  foot  and  hoof 

1,054 

765 

719 

1,340 

935 

819 

Fore  quarter,  right 

27,711 

19,194 

14,089 

46,507 

25,401 

21,605 

Hind  quarter,  right 

28,561 

18,582 

13,921 

47,873 

24,824 

20,820 

Left  half 

55,406 

36,500 

27,259 

95,368 

46,859 

44,904 

50 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  3. — Slaughter  House  Weights  of  Offal  Parts  (in  Grams). 


Steer 

505 

503 

532 

531 

504 

523 

525 

Age 

10  mo. 

11  mo. 

1 yr.  5 mo. 

1 yr.  6 mo. 

lyr.8mo. 

2yr.2mo. 

2yr.2mo. 

18  da. 

11  da. 

20  da. 

12  da. 

26  da. 

6 da. 

8 da. 

Group 

1 

2 

1 

3 

1 

2 

3 

Live  weight 

313,317 

270,566 

517,093 

212,466 

526,164 

380,880 

305,112 

Blood 

13,810 

13,058 

18,752 

9,457 

21,005 

15,287 

13,614 

Heart,  pericardium,  arteries 

2,106 

2,845 

4,843 

1,971 

3,065 

3,004 

1,320 

Heart,  marketable 

1,056 

1.251 

2,058 

1,011 

1,632 

1,477 

1,169 

Heart,  lean 

938 

1,063 

1,685 

791 

1,428 

1,188 

1,005 

Lungs  and  trachea 

2,498 

2,549 

3,870 

1,915 

3,628 

3,371 

1,658 

Brain 

376 

442 

406 

373 

423 

475 

467 

Spinal  cord 

161 

224 

237 

200 

301 

322 

244 

Tongue,  including  bones  and  larynx 

1,989 

1,365 

3,195 

1,728 

3,904 

3,702 

2,554 

Tongue,  marketable 

1,115 

789 

1,543 

894 

1,587 

1,678 

1,276 

Tongue  bones,  including  larynx 

190 

204 

340 

207 

367 

404 

252 

Gullet 

311 

442 

744 

494 

455 

803 

525 

Stomachs 

8,818 

5,765 

10,893 

5,343 

12,820 

9,342 

8,849 

5,943 

2,740 

1,272 

593 

Omasum  

2,208 

1,117 

Abomasum  

1,470 

893 

Intestines,  small 

| 4,852 

4,736  | 

3,792 

2,346 

| 9,526 

5,226 

5,960 

Intestines,  large 

2,132 

1,342 

Intestines  small;  length,  cm  

4,460 

4,039 

4,389 

2,926 

4,125 

3,797 

Intestines  large;  length,  cm  

909 

975 

762 

1,054 

866 

Neck  sweetbread 

425 

348 

321 

236 

396 

272 

150 

Heart  sweetbread 

337 

399 

120 

236 

532 

341 

145 

Spleen 

599 

661 

884 

481 

1,317 

847 

556 

Pancreas 

300 

289 

630 

297 

295 

300 

151 

Liver 

3,983 

3,646 

5,694 

2,205 

4,754 

3,268 

2,876 

Gall  bladder  and  gall 

215 

201 

284 

114 

697 

146 

126 

Gall  

185 

86 

Kidneys 

718 

655 

868 

506 

877 

703 

664 

Urinary  bladder 

80 

108 

308 

223 

254 

254 

193 

Penis 

251 

288 

302 

240 

310 

196 

157 

Diaphragm 

204 

917 

626 

270 

1,819 

362 

343 

Gaul  fat  

8,735 

686 

2,156 

1,132 

Stomach  fat 

6,344 

3,771 

4,940 

586 

12,272 

2,156 

1,180 

Intestinal  fat 

6,437 

3,614 

10,022 

1,627 

12,833 

3,603 

2,649 

Hide  and  hair 

23,026 

23,120 

33,988 

16,693 

41,144 

33,097 

27,813 

Dewclaws 

142 

112 

158 

90 

198 

115 

88 

Teeth 

268 

253 

494 

426 

338 

766 

690 

Horns  

342 

285 

228 

1,272 

1,167 

1,298 

Hoofs 

722 

661 

1,248 

700 

1,062 

948 

852 

Right  fore  foot  and  hoof 

1,218 

1,456 

1,937 

1,095 

1,767 

1,609 

1,380 

Left  fore  foot  and  hoof 

1,218 

1,469 

1,968 

1,096 

1,815 

1,664 

1,380 

Right  hind  foot  and  hoof 

1,251 

1,350 

1,932 

1,136 

1,736 

1,649 

1,320 

Left  hind  foot  and  hoof 

1,251 

1,383 

1,878 

1,137 

1,805 

1,608 

1,289 

Fore  quarter,  right 

45,336 

37,450 

79,605 

29,683 

76,580 

56,114 

43,204 

Hind  quarter,  right 

43,630 

36,954 

76,799 

29,429 

81,101 

53,785 

39,581 

Left  half 

87,885 

71,832 

156,225 

59,900 

148,728 

Studies  In  Animal  Nutrition — II 


51 


Table  4. — Slaughter  House  Weights  of  Offal  Parts  (in  Grams). 


Steer 

515 

507 

529 

527 

526 

524 

2yr.  9 mo. 

2 yr.  9 mo. 

3 yr.  2 mo. 

3 yr.  3 mo. 

3 yr.  4 mo. 

3 yr.  4 mo. 

Age 

19  days 

16  days 

21  days 

15  days 

13  days 

Group 

1 

2 

1 

1 

2 

3 

Live  weight 

743,361 

457,155 

690,704 

842,841 

479,846 

362,260 

Blood 

27,856 

20,316 

23,028 

27,382 

18,957 

17,019 

Heart,  pericardium,  arteries 

5,787 

4,278 

5,713 

8,891 

3,998 

2,953 

Heart,  marketable 

2,664 

2,061 

2,045 

3,246 

1,673 

1,585 

Heart,  lean 

1,890 

1,556 

1,419 

2,370 

1,246 

1,369 

Lungs  and  trachea 

3,653 

3,768 

4,272 

4,326 

3,797 

3,455 

Brain 

483 

482 

443 

407 

469 

508 

Spinal  cord 

245 

262 

261 

294 

191 

250 

Tongue,  including  bones  and  larynx 

4,229 

3,697 

4,021 

5,734 

3,626 

3,602 

Tongue,  marketable 

2,309 

1,791 

1,603 

1,904 

1,769 

1,666 

Tongue  bones,  including  larynx 

484 

467 

441 

420 

438 

435 

Gullet 

627 

709 

829 

1,090 

777 

1,340 

Stomachs 

13,171 

9,620 

12,101 

10,347 

10,985 

8,096 

6,007 

5,790 

6,091 

4,602 

1,069 

889 

896 

775 

3,179 

2,361 

2,460 

1,457 

Abomasum 

1,846 

1,307 

1,538 

1,262 

Intestines,  small 

j>  6,823 

2,985 

2,539 

2,257 

1,964 

Intestines,  large 

OylJ&O  \ 

2,304 

2,508 

1,877 

2,032 

Intestines, small; length,  cm 

j.  4 694 

q qaoJ 

3,818 

3,861 

3,515 

2,911 

Intestines, large;  length,  cm 

1,036 

1,072 

925 

747 

Neck  sweetbread 

484 

325 

514 

464 

269 

267 

Heart  sweetbread 

447 

318 

552 

573 

170 

274 

Spleen 

1,482 

1,132 

1,049 

1,226 

831 

757 

Pancreas 

424 

231 

824 

849 

498 

435 

Liver 

5,982 

3,788 

5,374 

5,720 

3,531 

3,019 

Gall  bladder  and  gall 

382 

130 

280 

149 

231 

296 

Gall 

190 

10 

143 

229 

Kidneys 

1,065 

752 

1,006 

1,244 

922 

766 

Urinary  bladder 

347 

282 

257 

295 

312 

361 

Penis 

225 

239 

436 

300 

250 

365 

Diaphragm 

1,107 

1,036 

645 

1,276 

807 

532 

Caul  fat 

9,263 

3,859 

13,993 

14,263 

2,763 

1,068 

Stomach  fat 

6,965 

3,097 

10,951 

9,669 

2,815 

1,244 

Intestinal  fat 

13,649 

4,351 

14,112 

24,585 

5,973 

2,695 

Hide  and  hair 

49,943 

34,473 

45,567 

46,240 

35,732 

30,092 

Dewclaws 

201 

152 

241 

240 

227 

194 

Teeth 

786 

712 

872 

782 

806 

Horns 

1,804 

1,600 

2,200 

1,266 

1,427 

1,227 

Hoofs 

1,692 

1,338 

1,934 

1,648 

1,300 

Hight  fore  foot  and  hoof 

2,346 

1,936 

2,213 

2,293 

2,031 

1,864 

Left  fore  foot  and  hoof 

2,312 

1,938 

2,230 

2,321 

1,984 

1,867 

Right  hind  foot  and  hoof 

2,473 

1,884 

2,331 

2,265 

1,862 

1,791 

Left  hind  foot  and  hoof 

2,607 

1,900 

2,362 

2,306 

1,863 

1,815 

Fore  quarter,  right 

115,806 

72,574 

111,434 

147,674 

75,917 

54,798 

Hind  quarter,  right 

112,738 

65,693 

112,704 

141,372 

70,858 

50,394 

Left  half 

229,567 

304,508 

52 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  5. — Slaughter  House  Weights  of  Offal  Parts  (in  Grams). 


Steer 

513 

502 

509 

501 

512 

500 

3 yr.  8 mo. 

3 yr.  8 mo. 

3 yr.  8 mo. 

3 yr. 

3yr.  11  mo. 

. 3yr.  11  mo. 

Age 

15  days 

19  days 

22  days 

11  mo. 

21  days 

26  days 

Group 

1 

2 

3 

1 

2 

3 

Live  weight 

854,650 

506,878 

439,814 

883,480 

548,050 

457,786 

Blood 

25,680 

19,728 

18,291 

28,710 

24,176 

21,269 

Heart,  per  cardium,  arteries 

7,427 

4,451 

3,645 

6,177 

5,158 

3,414 

Heart,  marketable 

2,311 

1,946 

1,660 

2,214 

1,955 

1,467 

Heart,  lean 

2,003 

1,680 

1,444 

1,882 

1,555 

1,284 

Lungs  and  trachea 

4,858 

3,696 

3,283 

3,838 

3,881 

3,747 

Brain 

413 

465 

389 

459 

466 

422 

Spinal  cord 

335 

335 

350 

298 

200 

410 

Tongue,  including  bones  and  larynx 

4,171 

4,107 

3,317 

5,294 

4,215 

3,541 

Tongue,  marketable 

1,942 

2,198 

1,555 

2,153 

1,766 

1,619 

Tongue  bones,  including  larynx 

455 

499 

425 

500 

526 

439 

Gullet 

1,152 

1,098 

1,081 

1,051 

973 

909 

Stomachs 

13,446 

10,832 

9,851 

14,185 

11,089 

10,995 

Rumen 

7,704 

7,216 

5,274 

8,769 

5,867 

6,446 

1,431 

1,064 

933 

1,068 

1 045 

Omasum 

2,951 

2,218 

2,337 

2,976 

2,643 

2,419 

Abomasum 

1,360 

1,398 

1,176 

1,507 

1,511 

1.085 

Intestines,  small 

3,543 

2,324 

2,186 

2,796 

3,067 

2,645 

Intestines,  large 

2,327 

1,829 

1,703 

2,079 

2,255 

1,890 

Intestines, small;  length,  cm 

4,480 

3,545 

3,255 

3,848 

4,481 

3,388 

Intestines, large;  length,  cm 

1,054 

970 

901 

1,001 

1,067 

945 

Neck  sweetbread 

581 

338 

267 

335 

235 

250 

Heart  sweetbread 

753 

164 

363 

449 

276 

288 

Spleen 

1,114 

921 

1,304 

1,178 

1,255 

1,054 

Pancreas 

873 

581 

562 

836 

736 

625 

Liver 

5,920 

3,716 

3,875 

6,161 

4,416 

4,634 

Gallbladder  and  gall 

127 

334 

225 

266 

300 

300 

Gall 

37 

241 

110 

176 

212 

241 

Kidneys 

1,015 

838 

774 

1,037 

1,074 

1,019 

Urinary  bladder 

227 

283 

253 

275 

366 

331 

Penis 

330 

305 

274 

339 

302 

271 

Diaphragm 

811 

561 

661 

779 

672 

692 

Caul  fat 

24.621 

2,738 

2,937 

14,688 

5,545 

3,930 

Stomach  fat 

7,860 

3,256 

2,240 

7,432 

3,658 

2,968 

Intestinal  fat 

21,290 

5,383 

4,745 

16,503 

8,251 

6.042 

Hide  and  hair 

45,286 

39,556 

37,614 

50,090 

41,268 

35,938 

Dewclaws 

308 

218 

196 

331 

234 

247 

Teeth 

874 

1,038 

838 

778 

710 

852 

Horns 

2,144 

1,949 

3,354 

1,810 

Hoofs 

1,872 

1,792 

1,394 

2,192 

1,490 

1,848 

Right  fore  foot  and  hoof 

2,452 

2,272 

1,941 

2,466 

2,210 

2,226 

Left  fore  foot  and  hoof 

2,426 

2,277 

1,904 

2,467 

2,184 

2,141 

Right  hind  foot  and  hoof 

2,283 

2,115 

1,828 

2,502 

2,043 

2,117 

Left  hind  foot  and  hoof 

2,432 

2,115 

1,875 

2,569 

2,008 

2,126 

Fore  quarter,  right 

146,424 

80,313 

70,406 

152.353 

90,208 

71,295 

Hind  quarter,  right 

132,093 

73,106 

63,182 

151,476 

79,228 

64,330 

Left  half | 

278,645 

152,147 

131,803 

305,356 

169,239 

136,107 

Studies  In  Animal  Nutrition— II 


53 


Table  6. — Slaughter  House  Weights  of  Carcass  Parts  (in  Grams). 


Steer 

556 

554 

555 

557 

552 

548 

Age 

3 months 

3 months 

3 months 

5 mo.  17  da. 

5 mo.  7 da. 

5 mo.  9 da 

Group 

[l 

2 

3 

1 

2 

3 

Head,  total 

3,338 

2,895 

2,833 

5,614 

4,059 

4,033 

Lean,  total 

702 

667 

765 

1,048 

795 

996 

Fat,  total 

132 

141 

143 

728 

308 

118 

Bone,  total 

2,504 

2,087 

1,925 

3,838 

2,956 

2,901 

Shin,  right 

1,765 

1,468 

1,637 

2,465 

1,700 

1,592 

Lean,  right 

840 

645 

829 

1,159 

864 

735 

Fat,  right 

81 

81 

71 

194 

70 

51 

Bone,  right 

851 

732 

727 

1,123 

775 

792 

Neck,  right 

517 

414 

502 

769 

428 

527 

Lean,  right 

344 

231 

287 

423 

266 

287 

71 

51 

149 

54 

55 

Bone,  right 

100 

133 

215 

206 

114 

191 

Chuck,  right 

7.600 

6,744 

5,863 

13,088 

7,851 

6,866 

Lean,  right 

5,465 

4,795 

4,298 

8,783 

5,613 

5,033 

Fat,  right 

323 

188 

116 

1,718 

409 

193 

Bone,  right 

1,767 

1,733 

1,448 

2,613 

1,840 

1,624 

Plate,  right 

3,048 

2,245 

2,107 

6,739 

2,940 

2,525 

Lean  right 

2,096 

1,475 

1,461 

3,710 

1,887 

1,655 

Fat,  right 

156 

125 

42 

1,769 

288 

131 

Bone,  right 

773 

628 

605 

1,241 

750 

741 

Rib  right 

2.485 

2,144 

1,915 

5,276 

3,120 

2,412 

Lean,  right 

1,611 

1,457 

1,256 

3,186 

1,995 

1,570 

Fat,  right 

39 

808 

57 

20 

Bone,  right 

823 

677 

628 

1,286 

1,042 

807 

Loin,  right 

5,029 

4,088 

3,102 

9,367 

5,147 

3,864 

Lean,  right 

3,469 

2,906 

2,309 

5,857 

3,517 

2,834 

Fat,  right 

410 

214 

137 

2,236 

521 

192 

Bone,  right 

1,134 

934 

632 

1,239 

1,096 

826 

Kidney,  Fat,  right 

210 

120 

65 

1,614 

250 

142 

Flank,  right 

868 

533 

549 

2,169 

935 

630 

Lean,  right 

599 

384 

453 

1,117 

597 

445 

Fat,  right 

236 

134 

78 

1,051 

330 

170 

Bone,  right 

16 

11 

13 

8 

8 

16 

Rump,  right 

926 

891 

703 

1,779 

987 

811 

Lean,  right 

548 

453 

401 

814 

551 

485 

Fat,  right 

74 

73 

37 

477 

112 

36 

Bone,  right 

304 

366 

253 

480 

320 

285 

Round,  right 

6,106 

5,064 

4,600 

10,090 

6,230 

5,811 

Lean,  right 

4,736 

3,959 

3,563 

7,480 

4,915 

4,604 

Fat,  right 

343 

260 

188 

1.266 

454 

260 

Bone,  right 

987 

851 

826 

1,343 

813 

949 

Shank,  right 

1,478 

1,395 

1,309 

2,064 

1,429 

1,305 

Lean,  right 

447 

400 

417 

628 

495 

415 

Fat,  right 

36 

26 

27 

117 

59 

30 

Bone,  right 

976 

959 

852 

1,305 

871 

856 

Tail,  total 

172 

200 

122 

332 

237 

147 

Lean,  total 

80 

75 

48 

106 

106 

53 

Fat,  total 

33 

Bone,  total 

92 

125 

74 

193 

131 

94 

54 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  7. — Slaughter  House  Weights  of  Carcass  Parts  (in  Grams). 


Steer 

547 

550 

558 

541 

538 

540 

Age 

8 mo.  5 da. 

8 mo.  14  da. 

8 mo.  12  da. 

10mo.22da. 

10mo.26da. 

11  mo.  2 da. 

Group 

1 

2 

3 

1 

2 

3 

Head,  total 

6,461 

5,108 

4,764 

6,939 

5,178 

4,747 

Lean,  total 

2,532 

1,704 

1,388 

1,388 

1,118 

890 

Fat,  total 

808 

384 

312 

460 

280 

344 

Bone,  total 

3,121 

3,020 

3,064 

5,091 

3,780 

3,513 

Shin,  right 

2,487 

1,951 

1,705 

3,712 

2,400 

2,228 

Lean,  right 

1,378 

991 

778 

1,927 

1,197 

1,167 

Fat,  right 

169 

65 

51 

318 

96 

69 

Bone,  right 

928 

876 

864 

1,473 

1,104 

976 

Neck,  right 

525 

451 

343 

1,500 

856 

689 

Lean  right 

303 

259 

209 

976 

503 

452 

Fat,  right 

115 

40 

20 

183 

164 

20 

Bone,  right 

109 

147 

103 

338 

200 

213 

Chuck,  right 

13,902 

9,962 

7,557 

22,782 

13,036 

10,763 

Lean,  right 

10,100 

6,880 

5,343 

16,871 

9,442 

7,771 

Fat,  right 

1,300 

728 

269 

2,210 

1,033 

746 

Bone,  right 

2,451 

2,331 

1,925 

3,310 

2,506 

2,235 

Plate,  right 

5,804 

3,745 

2,055 

10,165 

5,120 

4,374 

Lean  right 

3,574 

2,424 

1,386 

6,285 

3,197 

2,798 

Fat,  right 

1,240 

448 

136 

2,463 

829 

646 

Bone,  right 

969 

762 

542 

1,415 

1,062 

923 

Rib,  right 

4,932 

3,085 

2,380 

8,630 

3,886 

3,418 

Lean,  right 

3,280 

2,084 

1,593 

5,794 

2,598 

2,361 

Fat,  right 

551 

120 

28 

1,217 

213 

162 

Bone,  right 

1,086 

870 

758 

1,581 

1,077 

912 

Loin,  right 

10,373 

6,840 

4,359 

17,833 

8,957 

7,803 

Lean,  right 

6,788 

4,536 

2,981 

11,708 

6,366 

5,350 

Fat,  right 

1,986 

1,067 

300 

4,044 

1,180 

1,154 

Bone,  right 

1,566 

1,232 

1,069 

1,958 

1,348 

1,275 

Kidney,  Fat,  right 

815 

378 

113 

3,028 

311 

341 

Flank,  right 

2,317 

1,101 

575 

3,725 

1,434 

973 

Lean,  right 

1,372 

656 

462 

1,670 

961 

657 

Fat,  right 

927 

420 

112 

2,037 

531 

311 

Bone,  right 

10 

8 

4 

17 

10 

8 

Rump,  right 

1,531 

1,180 

866 

2,765 

1,436 

1,300 

Lean,  right 

740 

631 

463 

1,378 

780 

764 

Fat,  right 

388 

200 

52 

849 

268 

218 

Bone,  right 

380 

350 

339 

528 

366 

311 

Round,  right 

10,871 

7,391 

6,223 

17,065 

10,294 

8,305 

Lean,  right 

8,546 

5,860 

4,382 

13,500 

8,162 

6,728 

Fat,  right 

1,075 

472 

322 

1,927 

851 

405 

Bone'  right 

1,244 

1,050 

1,047 

1,625 

1,278 

1,175 

Shank,  right 

2,285 

1,502 

1,537 

3,042 

2,106 

1,810 

Lean,  right 

856 

435 

432 

981 

754 

629 

Fat,  right 

148 

56 

42 

189 

48 

16 

Bone,  right 

1,267 

1,011 

1,053 

1,872 

1,304 

1,152 

Tail,  total 

292 

186 

98 

381 

229 

217 

Lean,  total 

138 

88 

18 

160 

82 

36 

Fat,  total 

9 

4 

54 

16 

4 

Bone,  total 

144 

94 

80 

206 

135 

138 

Studies  In  Animal  Nutrition — II 


55 


Table  8. — Slaughter  House  Weights  of  Carcass  Parts  (in  Grams). 


Steer 

505 

503 

532 

531 

504 

523 

525 

Age 

10  mo. 

11  mo. 

1 yr.  5 mo. 

1 yr.  6 mo. 

lyr.8  mo. 

2yr.2  mo. 

2yr.2  mo 

18  da. 

11  da. 

20  da. 

12  da. 

26  da. 

6 da. 

8 da. 

Group  

1 

2 

1 

3 

1 

2 

3 

Head,  total 

6,977 

8,321 

10,510 

6,251 

11,035 

9,848 

8,536 

Lean,  total 

1,494 

1,728 

2,486 

1,310 

2,656 

2,500 

2,078 

Fat,  total 

814 

886 

1,406 

280 

648 

306 

508 

Bone,  total 

4,669 

5,707 

6,514 

4,487 

7,675 

6,832 

5,758 

Shin,  right 

3,703 

3,330 

6,742 

3,107 

6,123 

4,941 

4,328 

Lean,  right 

2,017 

1,604 

3,544 

1,602 

3,176 

2,671 

2,421 

Fat,  right 

205 

147 

468 

112 

751 

263 

199 

Bone,  right 

1,479 

1,568 

2,703 

1,393 

2,164 

1,995 

1,681 

Neck,  right 

1,044 

1,159 

1,476 

1,292 

1,536 

1,391 

1,552 

Lean,  right 

641 

548 

839 

930 

844 

943 

1,025 

Fat,  right 

135 

181 

277 

118 

233 

58 

117 

Bone,  right 

264 

430 

370 

257 

454 

378 

398 

Chuck,  right 

21,957 

18,821 

38,288 

15,303 

34,645 

28,898 

20,626 

Lean,  right 

15,877 

14,040 

27,000 

11,524 

23,092 

21,723 

15,605 

Fat,  right 

2,519 

1,198 

4,986 

879 

5,735 

2,436 

1,165 

Bone,  right 

3,497 

3,384 

6,137 

2,792 

5,134 

4,515 

3,827 

Plate,  right 

9,943 

7,580 

18,479 

5,483 

18,905 

11,995 

8,556 

Lean  right 

5,861 

4,398 

10,017 

3,585 

9,617 

7,704 

5,651 

Fat,  right 

2,586 

1,475 

5,657 

609 

6,963 

2,028 

1,174 

Bone,  right 

1,478 

1,676 

2,689 

1,248 

2,249 

1,914 

1,695 

Rib,  right 

8,607 

6,625 

14,483 

4,483 

15,286 

8,753 

8,134 

Lean,  right 

5,650 

4,466 

8,678 

3,137 

9,253 

6,016 

5,833 

Fat,  right 

1,320 

420 

3,097 

169 

3,385 

761 

332 

Bone,  right 

1,596 

1,691 

2,611 

1,140 

2,546 

1,922 

1,944 

Loin,  right 

15,622 

13,475 

27,366 

9,641 

29,121 

18,844 

13,241 

Lean,  right 

9,843 

8,603 

18,068 

7,039 

16,838 

12,917 

9,355 

Fat,  right 

3.779 

2,873 

7,477 

1,132 

9,170 

3,188 

1.879 

Bone,  right 

1,937 

1,917 

3,123 

1,417 

2,925 

2,661 

1,964 

Kidney,  Fat,  right 

2,877 

1,063 

5,867 

363 

5,700 

1,555 

629 

Flank,  right 

3,665 

2,330 

6,692 

1,253 

8,280 

3,795 

2,598 

Lean,  right 

1,912 

1,480 

2,934 

861 

3,778 

2,279 

1,704 

Fat,  right 

1,738 

792 

3,710 

372 

4,467 

1,481 

852 

Bone,  right 

20 

42 

50 

25 

37 

27 

32 

Rump,  right 

2,786 

2,146 

5,190 

1,783 

6,631 

3,627 

2,876 

Lean,  right 

1,300 

1,012 

2,570 

1,001 

2,884 

1,799 

1,589 

Fat,  right 

914 

591 

1,459 

252 

2,539 

910 

488 

Bone,  right 

553 

525 

1,141 

530 

1,214 

885 

771 

Round,  right 

15,380 

14,600 

25,006 

13,349 

25,995 

21,613 

16,843 

Lean,  right 

11,592 

11,499 

19,032 

10,748 

18,619 

16,950 

13,762 

Fat,  right 

2,071 

1,109 

3,032 

745 

4,909 

2,278 

981 

Bone,  right 

1,698 

1,928 

2,812 

1,820 

2,404 

2,315 

2,023 

Shank,  right 

2,885 

3,043 

4,682 

2,656 

4,817 

3,874 

3,024 

Lean,  right 

1,019 

991 

1,773 

971 

1,541 

1,324 

1,020 

Fat,  right 

162 

171 

219 

55 

772 

84 

152 

Bone,  right 

1,693 

1,874 

2,675 

1,606 

2,469 

2,436 

1,820 

Tail,  total 

520 

385 

675 

280 

691 

630 

468 

Lean,  total 

214 

150 

318 

120 

222 

256 

178 

Fat,  total 

58 

24 

50 

12 

64 

38 

24 

Bone,  total 

240 

190 

266 

148 

316 

316 

224 

56 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  9. — Slaughter  House  Weights  of  Carcass  Parts  (in  Grams). 


Steer 

515 

507 

529 

527 

526 

524 

2 yr.  9 mo. 

2 yr.  9 mo. 

3 yr.  2 mo. 

3 yr.  3 mo. 

3 yr.  4 mo. 

3 yr.  4 mo. 

19  da. 

16  da. 

21  da. 

15  da. 

13  da. 

Group 

1 

2 

1 

1 

2 

3 

Head,  total 

14,869 

11,874 

13,249 

12,838 

11,643 

10,915 

2,990 

2,370 

3,362 

2,426 

2,382 

2,484 

1,352 

1,246 

822 

740 

9,395 

8,152 

8,032 

8,395 

7,593 

Shin,  right 

8,370 

6,313 

8,275 

8,824 

6,789 

5,333 

3,822 

3,519 

4,371 

3,704 

2,631 

1,538 

258 

1,281 

248 

189 

3,009 

2,510 

3,170 

2,820 

2,475 

Neck,  right 

2,288 

1,611 

1,587 

1,832 

1,812 

1,473 

1,110 

998 

729 

1,047 

902 

667 

179 

603 

333 

38 

508 

417 

508 

425 

526 

Chuck,  right 

49,301 

36,460 

47,355 

62,001 

35,906 

28,236 

31,074 

27,311 

36,160 

25,478 

21,342 

11,216 

2,777 

18,728 

3,756 

911 

6.756 

6,184 

6,927 

6,496 

5,922 

Plate,  right 

34,875 

16,603 

35,901 

46,616 

18,103 

10,486 

Lean  right  

13,189 

10,064 

17,706 

10,740 

6,548 

Fat,  right  ....  

18,346 

3,390 

25,650 

4,110 

1,023 

Bnne  right  

3,124 

3,066 

3,019 

3,217 

2,895 

Rib,  right 

21,008 

11,635 

18,316 

28,401 

13,371 

9,216 

Lean,  right 

9,508 

7,894 

8,995 

12,930 

8,632 

6,403 

Fat,  right 

8,142 

1,216 

6,691 

12,139 

1,860 

169 

Bone,  right 

3,232 

2,525 

2,576 

3,273 

2,845 

2,655 

Loin,  right 

43,844 

23,374 

43,272 

55,188 

25,010 

16,656 

Lean  right  

20,810 

14,862 

25,070 

15,720 

12.100 

Fat  right  

19,162 

5,094 

26,362 

5,817 

1,222 

Bone  right  

3,892 

3,253 

3,570 

3,422 

3,293 

Kidney,  Fat,  right 

4,961 

2,188 

5,112 

9,482 

1,612 

383 

Flank,  right 

12,112 

4,697 

13,514 

16,167 

4,933 

2,407 

Lean  right  

3,281 

2,459 

4,823 

2,241 

1,563 

Fat,  right  . 

8,753 

2,152 

11,310 

2,671 

760 

Bone  right  . 

60 

73 

22 

77 

68 

Rump,  right 

10,005 

5,140 

9,344 

13,721 

5,748 

3,232 

Lean  right  

3,443 

2,521 

4,450 

2,804 

1,616 

Fat,  right  

4,462 

1,347 

7,560 

1,493 

399 

Bone,  right 

2,037 

1,268 

1,630 

1,419 

1,212 

Round,  right 

34,172 

25,274 

40.959 

39,746 

28,102 

23,013 

Lean,  right 

21,471 

19,651 

25,698 

22,307 

18,857 

Fat,  right  

9,529 

2,689 

10,733 

2,508 

1,263 

Bone,  right 

3,172 

2,932 

3,223 

3,176 

2,939 

Shank,  right 

6,92(2 

4,865 

* 

6,434 

4,865 

4,202 

Lean,  right 

1,873 

1,864 

2,190 

1,772 

1,422 

Fat,  right 

1,105 

289 

837 

98 

05 

Bone  right 

3,941 

2,665 

3,398 

2,986 

2,656 

Tail,  total 

882 

730 

857 

778 

700 

553 

Lean  total 

312 

292 

362 

328 

218 

Fat,  total 

132 

24 

48 

40 

24 

Bone  total 

354 

354 

370 

332 

290 

^Weighed  with  round. 


Studies  In  Animal  Nutrition — II 


57 


Table  10. — Slaughter  House  Weights  of  Carcass  Parts  (in  Grams). 


Steer 

513 

502 

509 

501 

512 

500 

Age 

3 yr.  8 mo. 
15  da. 

3 yr.  8 mo. 
19  da. 

3 yr.  8 mo. 
22  da. 

3yr.  11  mo. 

3yr.  11  mo. 
21  da. 

3yr.  11  mo. 
26  da. 

Group 

1 

2 

3 

1 

2 

3 

Head,  total 

13,287 

12,902 

10,642 

14,702 

13,233 

11,824 

Lean,  total 

3,690 

3,386 

2,556 

4,004 

3,266 

2,930 

Fat,  total 

1,294 

438 

264 

606 

706 

380 

Bone,  total 

7,865 

9,078 

7,822 

9,962 

9,139 

8,514 

Shin,  right 

8,748 

7,032 

6,523 

9,018 

7,437 

7,355 

Lean,  right 

4,639 

3,719 

3,769 

4,670 

3,928 

4,199 

Fat,  right 

1,039 

355 

238 

1,223 

461 

316 

Bone,  right 

3,060 

2,970 

2,523 

3,085 

3,037 

2,805 

Neck,  right 

1,944 

1,585 

1,635 

1,841 

1,600 

1,660 

Lean,  right 

987 

944 

925 

833 

800 

874 

Fat,  right 

616 

188 

171 

558 

175 

198 

Bone,  right 

357 

462 

537 

447 

620 

589 

Chuck,  right 

61,419 

41,503 

35,550 

61,734 

43,567 

35,199 

Lean,  right 

36,491 

29,998 

26,143 

35,518 

30,271 

26,178 

Fat,  right 

17,376 

4,242 

2,849 

18,586 

5,510 

2,209 

Bone,  right 

7,406 

7,084 

6,304 

7,442 

7,648 

6,636 

Plate,  right 

46,181 

16,244 

14,923 

51,977 

23,006 

16,798 

Lean  right 

18,269 

10,056 

9,499 

18,093 

11,581 

10,651 

Fat,  right 

24,127 

3,412 

2,896 

30,290 

7,474 

3,052 

Bone,  right 

3,623 

2,713 

2,512 

3,487 

3,827 

3,094 

Rib,  right 

27,830 

13,821 

11,725 

27,783 

14,527 

10,321 

Lean  right 

12,372 

9,128 

8,180 

10,417 

8,454 

6,801 

Fat,  right 

11,804 

1,669 

989 

14,161 

2,699 

902 

Bone,  right 

3,548 

2,960 

2,493 

3,194 

3,469 

2,596 

Loin,  right 

52,503 

26,154 

22,766 

63,056 

28,051 

22,159 

Lean,  right 

22,255 

17,552 

15,418 

22,998 

16,031 

14,846 

Fat,  right 

24,964 

4,572 

3,785 

35,679 

7,654 

3,415 

Bone,  right 

4,268 

3,933 

3,436 

4,307 

4,374 

3,886 

Kidney,  Fat,  right 

7,245 

1,458 

788 

9,772 

2,370 

1,216 

Flank,  right 

15,560 

4,390 

3,558 

18,768 

5,502 

4,586 

£ Lean,  right 

4,237 

2,331 

2,117 

4,544 

1,847 

2,805 

Fat,  right 

11,254 

1,998 

1,383 

14,146 

3,571 

1,697 

? Bone,  right 

96 

82 

50 

47 

67 

81 

Rump,  right 

11,288 

5,243 

5,142 

13,018 

6,870 

5,041 

Lean,  right 

3,609 

2,998 

2,631 

3,816 

3,054 

2,471 

i Fat,  right 

5,932 

1,052 

1,054 

7,297 

2,188 

1,058 

Bone,  right 

1,732 

1,181 

1,430 

1,841 

1,632 

1,494 

Round,  right 

38,655 

28,023 

25,862 

39,970 

30,725 

26,073 

Lean,  right 

25,391 

22,213 

20,188 

25,065 

21,704 

19,949 

Fat,  right 

9,554 

2,310 

2,553 

11,142 

4,970 

2,468 

Bone,  right 

3,522 

3,352 

3,010 

3,632 

3,913 

3,559 

Shank,  right 

6,245 

5,077 

4,639 

6,381 

5,071 

4,657 

Lean,  right 

1,813 

1,815 

1,708 

1,837 

1,717 

1,548 

Fat,  right 

1,408 

293 

176 

980 

247 

185 

Bone,  right 

3,029 

2,989 

2,749 

3,564 

3,078 

2,875 

Tail,  total 

675 

893 

774 

854 

875 

842 

Lean,  total 

304 

384 

324 

496 

366 

406 

Fat,  total 

102 

42 

68 

118 

74 

68 

Bone,  total 

297 

441 

386 

304 

416 

386 

58 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  11. — Distribution  of  Carcass  and  Offal  Parts. 


Steer 

556 

554 

555 

557 

552 

548 

Age 

3 mo. 

3 mo. 

3 mo. 

5 mo. 
17  da. 

5 mo. 
7 da. 

5 mo. 
9 da. 

Group 

1 

2 

3 

1 

2 

3 

Live  weight 

111,489 

87,456 

84,725 

204,934 

116,491 

99,255 

Warm  empty  weight 

98,133 

78,071 

71,078 

172,797 

99,349 

85,988 

Percent  empty  weight  to  live  weight 

88.020 

89.269 

83.893 

84.318 

85.285 

86.633 

Percent  carcass  to  live  weight 

54.188 

57.319 

53.626 

53.710 

52.998 

54.388 

Percent  carcass  to  empty  weight 

61.563 

64.210 

63.923 

63.700 

62.143 

62.780 

Percent  carcass  + offal  fat  to  live  weight.  . . . 

55.446 

58.055 

54.172 

57.008 

54.530 

54.232 

Percent  carcass  + offal  fat  to  empty  weight. . 

62.992 

65.034 

64.573 

67.610 

63.938 

63.762 

Percent  offal  fat  to  empty  weight 

1.429 

0.825 

0.650 

3.910 

1.796 

0.983 

Percent  hide  and  hair  to  empty  weight 

10.510 

9.479 

9.257 

8.160 

10.601 

9.720 

Percent  blood  to  empty  weight 

6.241 

5.353 

6.372 

5.181 

5.253 

5.353 

Percent  heart  market  to  empty  weight 

0.531 

0.429 

0.443 

0.537 

0.565 

0.497 

Percent  lungs  and  trachea  to  empty  weight. . 

1.295 

1.162 

1.318 

1.141 

1.103 

1.261 

Percent  brain  and  spinal  cord  to  empty  weight 

0.406 

0.506 

0.497 

0.319 

0.469 

0.612 

Percent  stomach  to  empty  weight 

2.013 

1.547 

1.882 

2.512 

1.951 

2.192 

Percent  intestines  to  empty  weight 

2.745 

2.660 

3.437 

2.370 

2.802 

2.932 

Cm.  intestines  per  kilo  empty  weight 

34.83 

37.17 

34.61 

22.03 

31.41 

39.03 

Percent  of  liver  to  empty  weight 

1.793 

1.494 

1.745 

1.838 

1.189 

1.284 

Percent  gall  bladder  and  gall  to  empty  weight 

0.025 

0.029 

0.011 

0.086 

0.055 

0.040 

Percent  of  kidneys  to  empty  weight 

0.344 

0.679 

0.618 

0.376 

0.318 

0.411 

Percent  spleen  to  empty  weight 

0.306 

0.259 

0.235 

0.281 

0.286 

0.250 

Percent  pancreas  to  empty  weight 

0.098 

0.097 

0.149 

0.130 

0.102 

0.099 

Table  12. — Distribution  of  Carcass  and  Offal  Parts. 


Steer 

547 

550 

558 

541 

538 

540 

Age 

8 mo. 
5 da. 

8 mo. 
14  da. 

8 mo. 
12  da. 

10  mo. 
22  da. 

10  mo. 
26  da. 

11  mo. 
2 da. 

Group 

1 

2 

3 

1 

2 

3 

Live  weight 

206,175 

147,202 

108,191 

323,836 

180,930 

158,131 

Warm  empty  weight 

171,448 

121,112 

89,999 

288,297 

158,911 

137,726 

Percent  empty  weight  to  live  weight 

83.157 

82.277 

83.185 

89.025 

87.830 

87.096 

Percent  carcass  to  live  weight 

54.167 

50.459 

51.085 

58.594 

53.658 

55.226 

Percent  carcass  to  empty  weight 

65.138 

61.328 

61.411 

65.817 

61.093 

63.408 

Percent  carcass  + offal  fat  to  live  weight.  . . . 

56.048 

52.232 

51.851 

61.786 

55.566 

56.685 

Percent  carcass  + offal  fat  to  empty  weight. . 

67.401 

63.483 

62.332 

69.403 

63.266 

65.083 

Percent  offal  fat  to  empty  weight 

2.262 

2.155 

0.921 

3.586 

2.172 

1.675 

Percent  hide  and  hair  to  empty  weight 

8.526 

8.620 

9.042 

9.219 

9.654 

9.435 

Percent  blood  to  empty  weight 

5.081 

5.846 

5.185 

4.325 

4.543 

5.059 

Percent  heart  market  to  empty  weight 

0.450 

0.459 

0.562 

0.377 

0.549 

0.489 

Percent  lungs  and  trachea  to  empty  weight . . 

1.090 

1.205 

1.234 

0.828 

1.148 

1.090 

Percent  brain  and  spinal  cord  to  empty  weight 

0.268 

0.366 

0.578 

0.197 

0.306 

0.372 

Percent  stomach  to  empty  weight 

3.027 

3.165 

2.833 

2.732 

2.845 

2.827 

Percent  intestines  to  empty  weight 

2.610 

2.992 

2.908 

1.819 

2.294 

2.366 

Cm.  intestines  per  kilo  empty  weight 

20.78 

27.87 

34.43 

14.08 

25.19 

23.83 

Percent  of  liver  to  empty  weight 

1.663 

1.645 

1.502 

1.329 

1.245 

1.157 

Percent  gall  bladder  and  gall  to  empty  weight 

0.080 

0.097 

0.031 

0.095 

0.096 

0.060 

Percent  of  kidneys  to  empty  weight 

0.262 

0.313 

0.353 

0.223 

0.306 

0.264 

Percent  spleen  to  empty  weight 

0.228 

0.240 

0.214 

0.207 

0.208 

0.240 

Percent  pancreas  to  empty  weight 

0.117 

0.165 

0.170 

0.135 

0.131 

0.131 

Studies  In  Animal  Nutrition — II 


59 


Table  13. — Distribution  of  Carcass  and  Offal  Parts. 


Steer 

505 

503 

532 

531 

504 

523 

525 

10  mo. 

11  mo. 

lyr. 

lyr. 

lyr. 

2yr. 

2 yr. 

18  da. 

11  da. 

5 mo. 

6 mo. 

8 mo. 

2 mo. 

2 mo. 

20  da. 

12  da. 

26  da. 

6 da. 

8 da. 

Group 

1 

2 

1 

3 

1 

2 

3 

Live  weight 

313,317 

270,566 

517,093 

212,466 

526,164 

380,880 

305,112 

Warm  empty  weight 

274,357 

236,429 

459,025 

192,005 

475,854 

337,803 

265,587 

Percent  empty  weight  to  live  weight 

87.565 

87.383 

88.770 

90.370 

90.438 

88.690 

87.046 

Percent  carcass  to  live  weight 

56.445 

54.048 

60.459 

56.015 

58.235 

57.708 

54.265 

Percent  carcass  to  empty  weight 

64.460 

61.852 

68.107 

61.984 

64.391 

65.067 

62.341 

Percent  carcass  +,  offal  fat  to  live  weight 

60.524 

56.778 

65.042 

57.379 

63.006 

59.786 

55.891 

Percent  carcass  + offal  fat  to  empty  weight 

69.119 

64.976 

73.270 

63.494 

69.667 

67.410 

64.209 

Percent  offal  fat  to  empty  weight 

4.659 

3.124 

5.162 

1.510 

5.276 

2.343 

1.868 

Percent  hide  and  hair  to  empty  weight 

8.393 

9.779 

7.404 

8.694 

8.646 

9.798 

10.472 

Percent  blood  to  empty  weight 

5.034 

5.523 

4.085 

4.925 

4.414 

4.525 

5.126 

Percent  heart  market  to  empty  weight 

0.385 

0.529 

0.448 

0.527 

0.343 

0.437 

0.440 

Percent  lungs  and  trachea  to  empty  weight 

0.910 

1.078 

0.843 

0.997 

0.762 

0.998 

0.624 

Percent  brain  and  spinal  co^d  to  empty  weight. . 

0.196 

0.282 

0.140 

• 0.298 

0.152 

0.236 

0.268 

Percent  stomach  to  empty  weight 

3.214 

2.439 

2.373 

2.783 

2.694 

2.766 

3.332 

Percent  intestines  to  empty  weight 

1.768 

2.003 

1.291 

1.921 

2.002 

1.547 

2.244 

Cm.  intestines  per  kilo  empty  weight  

16.26 

20.93 

11.69 

19.21 

10.88 

13.80 

Percent  of  liver  to  empty  weight 

1.452 

1.542 

1.240 

1.148 

0.999 

0.967 

1.083 

Percent  gall  bladder  and  gall  to  empty  weight. . . 

0.078 

0.085 

0.062 

0.059 

0.146 

0.043 

0.047 

Percent  of  kidneys  to  empty  weight 

0.262 

0.277 

0.189 

0.264 

0.184 

0.208 

0.250 

Percent  spleen  to  empty  weight 

0.218 

0.280 

0.193 

0.251 

0.277 

0.251 

0.209 

Percent  pancreas  to  empty  weight 

0.109 

0.122 

0.131 

0.155 

0.062 

0.089 

0.057 

Table  14. — Distribution  of  Carcass  and  Offal  Parts. 


Steer 

515 

507 

529 

527 

526 

524 

Age 

2 yr.  9 mo. 
19  da. 

2 yr.  9 mo. 
16  da. 

3 yr.  2 mo. 
21  da. 

3 yr.  3 mo. 
15  da. 

3 yr.  4 mo. 

3 yr.  4 mo. 
13  da. 

Group 

1 

2 

1 

1 

2 

3 

Live  weight 

743,361 

457,155 

690,704 

842,841 

479,846 

362,260 

Warm  empty  weight 

671,917 

418.896 

637,507 

786,005 

427,995 

322,234 

Percent  empty  weight  to  live  weight 

90.389 

91.631 

92.298 

93.257 

89.194 

88.951 

Percent  carcass  to  live  weight 

61.489 

60.490 

65.687 

70.423 

61.176 

58.075 

Percent  carcass  to  empty  weight 

68.028 

66.015 

71.169 

75.515 

68.587 

65.289 

Percent  carcass  + offal  fat  to  live  weight. . . . 

65.509 

62.964 

71.342 

76.179 

63.583 

59.458 

Percent  carcass  + offal  fat  to  empty  weight. . 

72.474 

68.714 

77.295 

81.688 

71.286 

66.843 

Percent  offal  fat  to  empty  weight 

4.447 

2.699 

6.126 

6.173 

2.699 

1.554 

Percent  hide  and  hair  to  empty  weight 

7.433 

8.229 

7.148 

5.883 

8.349 

9.339 

Peroent  blood  to  empty  weight 

4.161 

4.850 

3.612 

3.484 

4.429 

5.282 

Percent  heart  market  to  empty  weight 

0.396 

0.492 

0.321 

0.413 

0.391 

0.492 

Percent  lungs  and  trachea  to  empty  weight . 

0.544 

0.900 

0.670 

0.550 

0.887 

1.072 

Percent  brain  and  spinal  cord  to  empty  weight 

0.108 

0.178 

0.110 

0.089 

0.154 

0.235 

Percent  stomach  to  empty  weight 

1.960 

2.297 

1.898 

1.316 

2.567 

2.512 

Percent  intestines  to  empty  weight 

1.015 

1.343 

0.830 

0.642 

0.966 

1.240 

Cm.  intestines  per  kilo  empty  weight 

6.99 

9.46 

6.05 

6.28 

10.37 

11  35 

Percent  of  liver  to  empty  weight 

0.890 

0.904 

0.843 

0.728 

0.825 

0.937 

Percent  gall  bladder  and  gall  to  empty  weight 

0.057 

0.031 

0.044 

0.019 

0.054 

0.092 

Percent  of  kidneys  to  empty  weight 

0.159 

0.180 

0.158 

0.158 

0.215 

0.238 

Percent  spleen  to  empty  weight 

0.221 

0.270 

0.165 

0.156 

0.194 

0.235 

Percent  pancreas  to  empty  weight 

0.063 

0.055 

0.129 

0.108 

0.116 

0.135 

60 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  15. — Distribution  of  Carcass  and  Offal  Parts. 


Steer 

513 

502 

509 

501 

512 

500 

Age 

3 yr.  8 mo. 
15  da. 

3 yr.  8 mo. 
19  da. 

3 yr.  8 mo. 
22  da. 

3yr.  11  mo. 

3yr.  11  mo. 
21  da. 

3yr.  11  mo. 
26  da. 

Group 

1 

2 

3 

1 

2 

3 

Live  weight 

853,007 

506,878 

439,814 

883,480 

548,050 

457,786 

Warm  empty  weight 

771,142 

444,424 

391,461 

814,914 

493,877 

407,833 

Percent  empty  weight  to  live  weight 

90.403 

87.679 

89.006 

92.239 

90.115 

89.088 

Percent  carcass  to  live  weight 

65.317 

60.284 

60.342 

68.953 

61.796 

59.358 

Percent  carcass  to  empty  weight 

72.252 

68.756 

67.795 

74.755 

68.575 

66.628 

Percent  carcass  + offal  fat  to  live  weight. . . 

71.621 

62.528 

62.598 

73.325 

64.981 

62.185 

Percent  carcass  + offal  fat  to  empty  weight. 

79.224 

71.315 

70.330 

79.494 

72.109 

69.801 

Percent  offal  fat  to  empty  weight 

6.973 

2.560 

2.535 

4.740 

3.534 

3.173 

Percent  hide  and  hair  to  empty  weight 

5.873 

8.901 

9.609 

6.147 

8.356 

8.812 

Percent  blood  to  empty  weight 

3.330 

4.439 

4.672 

3.523 

4.895 

5.215 

Percent  heart  market  to  empty  weight 

0.300 

0.438 

0.424 

0.272 

0.396 

0.360 

Percent  lungs  and  trachea  to  empty  weight. . 

0.630 

0.832 

0.839 

0.471 

0.786 

0.919 

Percent  brain  and  spinal  cord  to  empty  weight 

0.097 

0.180 

0.189 

0.093 

0.135 

0.204 

Percent  stomach  to  empty  weight 

1.744 

2.438 

2.516 

1.741 

2.245 

2.696 

Percent  intestines  to  empty  weight 

0.761 

0.934 

0.994 

0.598 

1.078 

1.112 

Cm.  intestines  per  kilo  empty  weight 

7.18 

10.16 

10.62 

5.95 

11.23 

10.62 

Percent  of  liver  to  empty  weight 

0.768 

0.836 

0.990 

0.756 

0.894 

1.136 

Percent  gall  bladder  and  gall  to  empty  weight 

0.016 

0.075 

0.057 

0.033 

0.061 

0.074 

Percent  of  kidneys  to  empty  weight 

0.132 

0.189 

0.198 

0.127 

0.217 

0.250 

Percent  spleen  to  empty  weight 

0.144 

0.207 

0.333 

0.145 

0.254 

0.258 

Percent  pancreas  to  empty  weight 

0.113 

0.131 

0.144 

0.103 

0.149 

0.153 

Table  16. — Main  Divisions  of  Empty  Animal  and  Loss  on  Cooling. 


Steer 

556 

554 

555 

557 

552 

548 

Age 

3 mo. 

3 mo. 

3 mo. 

5 mo.  17  da. 

5 mo.  7 da. 

5 mo.  9 da. 

Group 

1 

2 

3 

1 

2 

3 

Weights  of  parts  in  grams 


Warm  empty  weight 

98,133 

78,071 

71,078 

172,797 

99,349 

85,988 

Carcass 

60,414 

50,129 

45,435 

110,071 

61,738 

53,983 

Offal  fat 

1,402 

644 

462 

6,757 

1,784 

845 

Hide  and  hair 

10,314 

7,400 

6,580 

14,100 

10,532 

8,358 

Head,  tail,  feet,  etc 

6,813 

6,203 

5,653 

10,591 

8,081 

7,463 

Blood 

6,124 

4,197 

4,529 

8,952 

5,219 

4,603 

Organs 

11,488 

8,419 

8,489 

19,831 

10,701 

9,645 

Loss  on  cooling 

1,422 

300 

23 

724 

716 

1,658 

Percent  of  parts  to  warm  empty  weight 


Carcass 

61.563 

64.210 

63.923 

63.700 

62.143 

62.780 

Offal  fat 

1.429 

0.825 

0.650 

3.910 

1.796 

0.983 

Hide  and  hair 

10.510 

9.479 

9.257 

8.160 

10.601 

9.720 

Head,  tail,  feet,  etc 

6.943 

7.945 

7.953 

6.129 

8.134 

8.679 

Blood 

6.241 

5.353 

6.372 

5.181 

5.253 

5.353 

Organs 

11.707 

10.784 

11.943 

11.476 

10.771 

11.217 

Loss  on  cooling 

1.449 

0.384 

0.032 

0.419 

0.721 

1.928 

Studies  In  Animal  Nutrition — II 


61 


Tables  17,  18,  19. — Main  Divisions  of  Empty  Animal  and  Loss  on?Cooling. 


Steer 

547 

550 

558 

541 

538 

__  540 

Age 

8 mo. 

8 mo. 

8 mo. 

10  mo. 

10  mo. 

11  mo 

5 da. 

14  da. 

12  da. 

22  da. 

26  da. 

2 da. 

Group 

1 

2 

3 

1 

2 

3 

Weights  of  i 

parts  in  grams 

Warm  empty  weight 

171,448 

121,112 

89,999 

288,297 

158,911 

137,726 

Carcass 

111,678 

74,276 

55,269 

189,748 

97,084 

87,329 

Offal  fat 

3,879 

2,610 

829 

10,337 

3,452 

2,307 

Hide  and  hair 

14,618 

10,440 

8,138 

26,576 

15,342 

12,994 

Head,  tail,  feet,  etc 

11,802 

8,969 

8,289 

13,781 

9,921 

9,005 

Blood 

8,711 

7,080 

4,666 

12,470 

7,219 

6,967 

Organs 

19,822 

15,084 

11,128 

28,319 

17,879 

15,662 

Loss  on  cooling 

72 

1,377 

929 

8,054 

4,648 

5,941 

Percent  of  parts  to 

warm  empt 

y weight 

Carcass 

65.138 

61.328 

61.411 

65.817 

61.093 

63.408 

Offal  fat 

2.262 

2.155 

0.921 

3.586 

2.172 

1.675 

Hide  and  hair 

8.526 

8.620 

9.042 

9.219 

9.654 

9.435 

Head,  tail,  feet,  etc 

6.884 

7.406 

9.210 

4.780 

6.243 

6.538 

Blood 

5.081 

5.846 

5.185 

4.325 

4.543 

5.059 

Organs 

11.562 

12.455 

12.365 

9.823 

11.251 

11.372 

Loss  on  cooling 

0.042 

1.137 

1.032 

2.794 

2.925 

4.314 

Steer 

505 

503 

532 

531 

504 

523 

525 

Age 

10  mo. 

11  mo. 

1 yr.  5 mo. 

1 yr.  6 mo. 

1 yr.  8 mo. 

2 yr.  2 mo. 

2 yr.  2 mo . 

18  da. 

11  da. 

20  da. 

12  da. 

26  da. 

6 da. 

8 da. 

Group 

1 

2 

1 

3 

1 

2 

3 

Weights  of  parts  in  grams 


Warm  empty  weight 

274,357 

236,429 

459,025 

192,005 

475,854 

337,803 

265,587 

Carcass 

176,851 

146,236 

312,629 

119,012 

306,409 

219,798 

165,570 

Offal  fat 

12,781 

7,385 

23,697 

2,899 

25,105 

7,915 

4,961 

Hide  and  hair 

23,026 

23,120 

33,988 

16,693 

41,144 

33,097 

27,813 

Head,  tail,  feet.,  etc 

13,377 

15,218 

20,120 

11,718 

21,024 

23,460 

16,701 

Blood 

13,810 

13,058 

18,752 

9,457 

21,005 

15,287 

13,614 

Organs 

28,033 

25,676 

39,809 

20,313 

45,006 

32,570 

26,676 

Loss  on  cooling 

5,398 

3,164 

10,030 

12,701 

7,208 

5,676 

10,252 

Percent  of  parts  to 

warm  empt 

y weight 

Carcass 

64.460 

61.852 

68.107 

61.984 

64.391 

65.067 

62.341 

Offal  fat 

4.659 

3.124 

5.162 

1.510 

5.276 

2.343 

1.868 

Hide  and  hair 

8.393 

9.779 

7.404 

8.694 

8.646 

9.798 

10.472 

Head,  tail,  feet,  etc 

4.876 

6.437 

4.383 

6.103 

4.418 

6.945 

6.288 

Blood 

5.034 

5.523 

4.085 

4.925 

4.414 

4.525 

5.126 

Organs 

10.218 

10.860 

8.673 

10.579 

9.458 

9.642 

10.044 

Loss  on  cooling 

1.968 

1.338 

2.185 

6.615 

1.515 

1.680 

3.860 

Steer 

515 

507 

529 

527 

526 

524 

Age 

2 yr.  9 mo. 

2 yr.  9 mo. 

3 yr.  2 mo. 

3 yr.  3 mo. 

3 yr.  4 mo. 

3 yr.  4 ma 

19  da. 

16  da. 

21  da. 

15  da. 

13  da. 

Group 

1 

2 

1 

1 

2 

3 

Weights  of  parts  in  grams 

Warm  empty  weight 

671,917 

418,896 

637,507 

786,005 

427,995 

322,234 

Carcass 

457,088 

276,534 

453,705 

593,554 

293,550 

210,384 

Offal  fat 

29,877 

11,307 

39,056 

48,517 

11,551 

5,007 

Hide  and  hair 

49,943 

34,473 

45,567 

46,240 

35,732 

30,092 

Head,  tail,  feet,  etc 

28,764 

23,193 

26,124 

25,599 

22,957 

21,467 

Blood 

27,856 

20,316 

23,028 

27,382 

18,957 

17,019 

Organs 

51,173 

36,207 

43,425 

47,812 

35,360 

30,837 

Loss  on  cooling 

27,216 

16,866 

12,031 

12,363 

9,888 

7,428 

Percent  of  parts  to  warm  empty 


Carcass 

Offal  fat 

Hide  and  hair 

Head,  tail, feet,  etc 

Blood 

Organs 


68.028 

4.447 

7.433 

4.281 

4.161 

7.616 


66.015 

2.699 

8.229 

5.537 

4.850 

8.643 

4 n9« 


weight 

71.169 

6.126 

7.148 

4.098 

3.612 

6.812 

1 


75.515 

6.173 

5.883 

3.257 

3.484 

6.083 

1 K73 


68.587 
2.699 
8.349 
5.364 
4.429 
8.262 
o Qin 


65.289 

1.554 

9.339 

6.662 

5.282 

9.570 


62 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  20. — Main  Divisions  of  Empty  Animal  and  Loss  on  Cooling. 


Steer 

513 

502 

509 

501 

512 

500 

Age 

3 yr.  8 mo. 
15  da. 

3 yr.  8 mo. 
19  da. 

3 yr.  8 mo. 
22  da. 

3 yr.ll  mo. 

3 yr.ll  mo. 
21  da. 

3 yr.ll  mo. 
26  da. 

Group 

1 

2 

3 

1 

2 

3 

Weights  of  parts  in  grams 


Warm  empty  weight 

771,142 

444,424 

391,461 

814,914 

493,877 

407,833 

Carcass 

557,162 

305,566 

265,391 

609,185 

338,675 

271,732 

Offal  fat 

53,771 

11,377 

9,922 

38,625 

17,454 

12,940 

Hide  and  hair 

45,286 

39,556 

37,614 

50,090 

41,268 

35,938 

Head,  tail,  feet,  etc 

27,336 

26,278 

20,423 

30.523 

25,833 

22,814 

Blood 

25,680 

19,728 

18,291 

28,710 

24,176 

21,269 

Organs 

48,968 

36,679 

33,938 

47,332 

40,410 

36,998 

Loss  on  cooling 

13,067 

3,968 

4,097 

11,976 

5,864 

6,624 

Percent  of  parts  to  warm  empty  weight 


Carcass 

72.252 

68.756 

67.795 

74.755 

68.575 

66.628 

Offal  fat 

6.973 

2.560 

2.535 

4.740 

3.534 

3.173 

Hide  and  hair 

5.873 

8.901 

9.609 

6.147 

8.356 

8.812 

Head  tail  feet  etc 

3.545 

5.913 

5.217 

3.746 

5.231 

5.594 

5.215 

Blood 

3.330 

4.439 

4.672 

3.523 

4.895 

Organs  

6.350 

8.253 

8.670 

5.808 

8.182 

9.072 

1.624 

Loss  on  cooling  

1.694 

0.893 

1.047 

1.470 

1.187 

Table  21. — Proportion  of  Cuts  to  Empty  Weight  and  to  Carcass. 


Steer 

556 

554 

555 

557 

552 

548 

Age 

3 mo. 

3 mo. 

3 mo. 

5 mo. 
17  da. 

5 mo. 
7 da. 

5 mo. 
9 da. 

Group 

1 

2 

3 

1 

2 

3 

Warm  empty  weight 

98,133 

78,071 

71,078 

172,797 

99,349 

85,988 

Weight  of  carcass 

60,570 

50,908 

45,342 

111,842 

62,316 

53,416 

Percent  forequarters  to  empty  weight 

31.459 

33.549 

33.977 

32.967 

32.256 

32.432 

Percent  forequarters  to  carcass 

50.969 

51.450 

53.262 

50.934 

51.425 

52.209 

Percent  hindquarters  to  empty  weight 

30.263 

31.658 

29.815 

31.757 

30.468 

29.688 

Percent  hindquarters  to  carcass 

49.031 

48.550 

46.738 

49.065 

48.575 

47.791 

Percent  shins  to  empty  weight 

3.597 

3.761 

4.606 

2.853 

3.422 

3.703 

Percent  shins  to  carcass 

5.828 

5.767 

7.221 

4.408 

5.456 

5.961 

Percent  of  neck  to  empty  weight 

1.054 

1.061 

1.413 

0.890 

0.862 

1.226 

Percent  of  neck  to  carcass 

1.707 

1.626 

1.214 

1.375 

1.374 

1.973 

Percent  chucks  to  empty  weight 

15.489 

17.277 

16.497 

15.148 

15.805 

15.970 

Percent  chucks  to  carcass 

25.095 

26.489 

25.861 

23.404 

25.197 

25.708 

Percent  of  plates  to  empty  weight 

6.212 

5.751 

5.929 

7.800 

5.919 

5.873 

Percent  of  plates  to  carcass 

10.064 

8.820 

9.294 

12.051 

9.436 

9.454 

Percent  ribs  to  empty  weight 

5.065 

5.492 

5.388 

6.107 

6.281 

5.610 

Percent  ribs  to  carcass 

8.205 

8.423 

8.447 

9.435 

10.013 

9.031 

Percent  of  loin  to  to  empty  weight 

10.249 

10.473 

8.728 

10.842 

10.361 

8.987 

Percent  loin  to  carcass 

16.606 

16.060 

13.683 

16.750 

16.519 

14.468 

Percent  kidney  fat  + kidney  to  empty  weight 

0.772 

0.986 

0.801 

2.244 

0.821 

0.741 

Percent  kidney  fat  + kidney  to  carcass 

1.251 

1.512 

1.255 

3.466 

1.309 

1.193 

Percent  flanks  to  empty  weight 

1.769 

1.365 

1.432 

2.510 

1.882 

1.465 

Percent  flanks  to  carcass 

2.866 

2.094 

2.245 

3.879 

3.001 

2.359 

Percent  rumps  to  empty  weight 

1.887 

2.283 

1.978 

2.059 

1.987 

1.886 

Percent  rumps  to  carcass 

3.058 

3.500 

3.101 

3.181 

3.168 

3.037 

Percent  rounds  to  empty  weight 

12.444 

12.973 

12.944 

11.678 

12.542 

13.516 

Percent  rounds  to  carcass 

20.162 

19.893 

20.290 

18.043 

19.995 

21.758 

Percent  shanks  to  empty  weight 

3.012 

3.574 

3.683 

2.390 

2.877 

3.035 

Percent  shanks  to  carcass 

4.880 

5.480 

5.774 

3.691 

4.586 

4.886 

Percent  head  to  empty  weight 

3.402 

3.708 

3.986 

3.249 

4.086 

4.690 

Percent  tail  to  empty  weight 

0.175 

0.256 

0.172 

0.192 

0.239 

0.171 

Studies  In  Animal  Nutrition — II 


63 


Table  22. — Proportion  of  Cuts  to  Empty  Weight  and  to  Carcass. 


Steer 

547 

550 

558 

541 

538 

540 

Age 

8 mo. 
5 da. 

8 mo. 
14  da. 

8 mo. 
12  da. 

10  mo. 
22  da. 

10  mo. 
26  da. 

11  mo. 
2 da. 

Group 

1 

2 

3 

1 

2 

3 

Warm  empty  weight 

171,448 

121,112 

89,999 

288,297 

158,911 

137,726 

Weight  of  carcass 

112,544 

75,552 

56.020 

188,760 

100,450 

84,850 

Percent  forequarters  to  empty  weight 

32.326 

31.696 

31.309 

32.263 

31.969 

31.374 

Percent  forequarters  to  carcass 

49.245 

50.810 

50.300 

49.276 

50.574 

50.925 

Percent  hindquarters  to  empty  weight 

33.317 

30.686 

30.963 

33.211 

31.243 

30.234 

Percent  hindquarters  to  carcass 

50.755 

49.190 

49.700 

50.724 

49.426 

49.075 

Percent  shins  to  empty  weight 

2.901 

3.222 

3.789 

2.575 

3.021 

3.235 

Percent  shins  to  carcass 

4.420 

5.165 

6.087 

3.933 

4.778 

5.252 

Percent  of  neck  to  empty  weight 

0.612 

0.745 

0.762 

1.041 

1.077 

1.001 

Percent  of  neck  to  carcass 

.933 

1.194 

1.225 

1.589 

1.704 

1.624 

Percent  chucks  to  empty  weight 

16.217 

16.451 

16.794 

15.805 

16.407 

15.630 

Percent  chucks  to  carcass 

24.705 

26.371 

26.980 

24.139 

25.955 

25.369 

Percent  of  plates  to  empty  weight 

6.771 

6.184 

4.567 

7.052 

6.444 

6.352 

Percent  of  plates  to  carcass 

10.314 

9.914 

7.337 

10.770 

10.194 

10.310 

Percent  ribs  to  empty  weight 

5.753 

5.094 

5.289 

5.987 

4.891 

4.964 

Percent  ribs  to  carcass 

8.766 

8.167 

8.497 

9.144 

7.737 

8.057 

Percent  of  loin  to  empty  weight 

12.100 

11.295 

9.687 

12.371 

11.273 

11.331 

Percent  loin  to  carcass 

18.434 

18.107 

15.562 

18.895 

17.834 

18.392 

Percent  kidney  fat  + kidney  to  empty  weight 

1.213 

0.937 

0.604 

2.324 

0.698 

0.759 

Percent  kidney  fat  + kidney  to  carcass 

1.848 

1.502 

0.971 

3.550 

1.104 

1.232 

Percent  flanks  to  empty  weight 

2.703 

1.818 

1.278 

2.584 

1.805 

1.413 

Percent  flanks  to  carcass 

4.118 

2.915 

2.053 

3.947 

2.855 

2.293 

Percent  rumps  to  empty  weight 

1.786 

1.949 

1.924 

1.918 

1.807 

1.888 

Percent  rumps  to  carcass 

2.721 

3.124 

3.092 

2.930 

2.859 

3.064 

Percent  rounds  to  empty  weight 

12.681 

12.205 

13.829 

11.838 

12.956 

12.060 

Percent  rounds  to  carcass 

19.319 

19.565 

22.217 

18.081 

20.496 

19.576 

Percent  shanks  to  empty  weight 

2.666 

2.480 

3.416 

2.110 

2.651 

2.628 

Percent  shanks  to  carcass 

4.061 

3.976 

5.487 

3.223 

4.193 

4.266 

Percent  head  to  empty  weight 

3.768 

4.218 

5.293 

2.407 

3.260 

3.447 

Percent  tail  to  empty  weight 

0.170 

0.154 

0.109 

0.132 

0.144 

0.158 

Table  23. — Proportion  of  Cuts  to  Empty  Weight  and  to  Carcass. 


Steer 

505 

503 

532 

531 

504 

523 

525 

10  mo. 
18  da. 

11  mo. 
11  da. 

1 yr. 
5 mo. 
20  da. 

1 yr. 
6 mo. 
12  da. 

1 yr. 
8 mo. 
26  da. 

2 yr. 
2 mo. 
' 6 da. 

2 yr. 
2 mo. 
8 da. 

Group 

1 

2 

1 

3 

1 

2 

3 

Warm  empty  weight 

274,357 

236,429 

459,025 

192,005 

475,854 

337,803 

265,587 

Weight  of  carcass 

177,932 

148,805 

312,808 

118,224 

315,362 

219,798 

165,570 

Percent  forequarters  to  empty  weight 

33.049 

31.680 

34.684 

30.919 

32.187 

33.223 

32.535 

Percent  foreauarters  to  carcass 

50.959 

50.334 

50.897 

50.215 

48.566 

51.060 

52.188 

Percent  hindquarters  to  empty  weight 

31.805 

31.260 

33.462 

30.654 

34.087 

31.844 

29.806 

Percent  hindquarters  to  carcass 

49.041 

49.688 

49.103 

49.785 

51.434 

48.940 

47.812 

Percent  shins  to  empty  weight 

2.699 

2.817 

2.938 

3.236 

2.574 

i 2 . 925 

3.259 

Percent  shins  to  carcass 

4.162 

4.476 

4.311 

5.256 

3.883 

4.496 

5.228 

Percent  of  neck  to  empty  weight 

0.761 

0.980 

0.643 

1.346 

0.646 

0.824 

1.169 

Percent  of  neck  to  carcass 

1.173 

1.558 

0.944 

2.186 

0.974 

M.266 

1.875 

Percent  chucks  to  empty  weight 

16.006 

15.921 

16.682 

15.940 

14.561 

17.109 

15.532 

Percent  chucks  to  carcass 

24.680 

25.296 

24.480 

25.888 

21.972 

26.295 

24.915 

Percent  of  plates  to  empty  weight 

7.248 

6.412 

8.051 

5.711 

7.946 

7.102 

6.443 

Percent  of  plates  to  carcass 

11.176 

10.188 

11.815 

9.276 

11.990 

10.915 

10.335 

Percent  ribs  to  empty  weight 

6.274 

5.604 

6.310 

4.670 

6.424 

{ 5.182 

i 6. 125 

Percent  ribs  to  carcass 

9.674 

8.904 

9.260 

7.584 

9.694 

17.965 

9.825 

Percent  of  loin  to  empty  weight 

11.388 

11.399 

11.924 

10.042 

12.239 

11.156 

9.971 

Percent  loin  to  carcass 

17.560 

18.111 

17.497 

16.310 

18.468 

17.147 

15.994 

Percent  kidney  fat  + kidney  to  empty  weight. . 

2.360 

1.176 

2.745 

0.642 

2.580 

1.129 

0.724 

Percent  kidney  fat  -|-  kidney  to  carcass 

3.637 

1.869 

4.029 

1.042 

3.893 

1.735 

1.161 

Percent  flanks  to  empty  weight 

2.672 

1.971 

2.916 

1.305 

3.480 

2.247 

1.956 

Percent  flanks  to  carcass 

4.120 

3 . 132 

4.279 

2.120 

5.251 

3.453 

3.138 

Percent  rumps  to  empty  weight 

2.031 

1.815 

2.261 

1.857 

2.787 

2.147 

2.166 

Percent  rumps  to  carcass 

3.132 

2.884 

3.318 

3.016 

4.205 

3.300 

3.474 

Percent  rounds  to  empty  weight 

11.212 

12.350 

10  895 

13.905 

10.926 

12.796 

12.684 

Percent  rounds  to  carcass 

17.288 

19.623 

15.988 

22.583 

16.486 

19.666 

20.345 

Percent  ehanks  to  empty  weight 

2.103 

2.574 

2 040 

2.767 

2.025 

2.294 

2.277 

Percent  shanks  to  carcass 

3.243 

4.090 

2.994 

4.493 

3.055 

3.525 

3.653 

Percent  head  to  empty  weight 

2.543 

3.519 

2 290 

3.256 

2.319 

2.915 

3.214 

Percent  tail  to  empty  weight 

0.190 

0.163 

0.147 

0.146 

0.145 

0.186 

0.176 

64 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  24. — Proportion  of  Cuts  to  Empty  Weight  and  to  Carcass. 


Steer 

515 

507 

529 

527 

526 

524 

2 yr.  9 mo. 

2 yr.  9 mo. 

3 yr.  2 mo. 

3 yr.  3 mo. 

3 yr.  4 mo. 

3 yr.  4 mo. 

Age 

19  da. 

16  da. 

21  da. 

15  da. 

13  da. 

Group 

1 

2 

1 

1 

2 

3 

Warm  empty  weight 

671,917 

418,896 

637,507 

786,005 

427,995 

322,234 

Wt.  of  carcass 

457,088 

276,534 

448,276 

578,092 

293,550 

210,384 

Percent  forequarters  to  empty  weight 

34.470 

34.650 

34.959 

37.576 

35.476 

34.011 

Percent  forequarters  to  carcass 

50.671 

52.488 

49.717 

51.090 

51.723 

52.093 

Percent  hindquarters  to  empty  weight 

33.557 

31.365 

35.358 

35.972 

33.112 

31.278 

Percent  hindquarters  to  carcass 

49.329 

47.512 

50.283 

48.910 

48.277 

47.907 

Percent  shins  to  empty  weight 

2.491 

3.014 

2.596 

2.245 

3.172 

3.310 

Percent  shins  to  carcass 

3.662 

4.566 

3.692 

3.053 

4.625 

5.070 

Percent  of  neck  to  empty  weight 

0.681 

0.769 

0.498 

0.466 

0.847 

0.914 

Percent  of  neck  to  carcass 

1.001 

1.165 

0.708 

0.634 

1.235 

1.400 

Percent  chucks  to  empty  weight 

14.675 

17.408 

14.856 

15.776 

16.779 

17.525 

Percent  chucks  to  carcass 

21.572 

26.369 

21.128 

21.450 

24.463 

26.842 

Percent  of  plates  to  empty  weight 

10.381 

7.927 

11.263 

11.862 

8.459 

6.508 

Percent  of  plates  to  carcass 

15.260 

12.008 

16.017 

16.128 

12.334 

9.968 

Percent  ribs  to  empty  weight 

6.253 

5.555 

5.746 

7.227 

6.248 

5.720 

Percent  of  loin  to  empty  weight 

9.192 

8.415 

8.172 

9.826 

9.110 

8.761 

Percent  ribs  to  carcass 

13.050 

11.160 

13.575 

14.043 

11.687 

10.338 

Percent  loin  to  carcass 

19.184 

16.905 

19.306 

19.093 

17.040 

15.834 

Percent  kidney  fat  + kidney  to  empty  weight 

1.635 

1.224 

1.762 

2.571 

0.969 

1.762 

Percent  kidney  fat  + kidney  to  carcass 

2.404 

1.854 

2.505 

3.496 

1.412 

2.699 

Percent  flanks  to  empty  weight 

3.605 

2.243 

4.240 

4.114 

2.305 

1.494 

Percent  flanks  to  carcass 

5.300 

3.397 

6.029 

5.593 

3.361 

2.288 

Percent  rumps  to  empty  weight 

2.978 

2.454 

2.931 

3.491 

2.686 

2.006 

Percent  rumps  to  carcass 

4.378 

3.717 

4.169 

4.747 

3.916 

3.072 

Percent  rounds  to  empty  weight 

10.171 

12.067 

12.850 

10.113 

13.132 

14.283 

Percent  rounds  to  carcass 

14.952 

18.279 

18.274 

13.751 

19.146 

21.877 

Percent  shanks  to  empty  weight  

2 060 

2 323 

1 637 

2.273 

2.608 

Percent  shanks  to  carcass 

3.029 

3.519 

2.226 

3.315 

3.995 

Percent  head  to  empty  weight 

2.213 

2.835 

2.078 

1.633 

2.720 

3.387 

Percent  tail  to  empty  weight 

0.131 

0.174 

0.134 

0.099 

0.164 

0.172 

Table  25. — Proportion  of  Cuts  to  Empty  Weight  and  to  Carcass. 


Steer ' 

513 

502 

509 

501 

512 

500 

Age 

3 yr.  8 mo. 
15  da. 

3 yr.  8 mo. 
19  da. 

3 yr.  8 mo. 
22  da. 

3yr.  11  mo. 

3yr.  11  mo. 
21  da. 

3yr.  11  mo. 
26  da. 

Group 

1 

2 

3 

1 

2 

3 

Warm  empty  weight 

771,142 

444,424 

391,461 

814,914 

493,877 

407,833 

Weight  of  carcass 

557,034 

306,838 

267,176 

607,658 

338,872 

371,250 

Percent  forequarters  to  empty  weight 

37.976 

36.143 

35.971 

37.391 

36.531 

34.963 

Percent  forequarters  to  carcass 

52.573 

52.349 

52.704 

50.144 

53.240 

52.568 

Percent  hindquarters  to  empty  weight 

34.259 

32.899 

32.280 

37.176 

32.084 

31.547 

Percent  hindquarters  to  carcass 

47.427 

47.651 

47.296 

49.856 

46.760 

47.432 

Percent  shins  to  empty  weight 

2.269 

3.165 

3.333 

2.213 

3.012 

3.607 

Percent  shins  to  carcass 

3.141 

4.584 

4.883 

2.968 

4.389 

5.423 

Percent  of  neck  to  empty  weight 

0.504 

0.713 

0.835 

0.452 

0.648 

0.814 

Percent  of  neck  to  carcass 

0.698 

1.033 

1.224 

0.606 

0.944 

1.224 

Percent  chucks  to  empty  weight 

15.929 

18.677 

18.163 

15.151 

17.643 

17.261 

Percent  chucks  to  carcass 

22.052 

27.052 

26.612 

20.319 

25.713 

25.953 

Percent  of  plates  to  empty  weight 

11.977 

7.310 

7.624 

12.756 

9.316 

8.238 

Percent  of  plates  to  carcass 

16.581 

10.588 

11.171 

17.107 

13.578 

12.386 

Percent  ribs  to  empty  weight 

7.218 

6.220 

5.990 

6.819 

5.883 

5.061 

Percent  ribs  to  carcass 

9.992 

9.009 

8.777 

9.144 

8.574 

7.610 

Percent  of  loin  to  empty  weight 

13.617 

11.770 

11.631 

15.476 

11.360 

10.867 

Percent  loin  to  carcass 

18.851 

17.047 

17.042 

20.754 

16.556 

16.338 

Percent  kidney  fat  + kidney  to  empty  weight 

2.011 

0.845 

0.600 

2.526 

1.177 

0.846 

Percent  kidney  fat  + kidney  to  carcass 

2.783 

1.223 

0.880 

3.387 

1.716 

1.272 

Percent  flanks  to  empty  weight 

4.036 

1.976 

1.818 

4.606 

2.228 

2.249 

Percent  flanks  to  carcass 

5.587 

2.861 

2.663 

6.177 

3.247 

3.381 

Percent  rumps  to  empty  weight 

2.928 

2.359 

2.627 

3.195 

2.782 

2.472 

Percent  rumps  to  carcass 

4.053 

3.417 

3.849 

4.285 

4.055 

3.717 

Percent  rounds  to  empty  weight 

10.025 

12.611 

13.213 

9.810 

12.442 

12.786 

Percent  rounds  to  carcass 

13.879 

18.266 

19.360 

13.139 

18.134 

19.224 

Percent  shanks  to  empty  weight 

1.620 

2.285 

2.370 

1.566 

2.054 

2.284 

Percent  shanks  to  carcass 

2.242 

3.309 

3.473 

2.100 

2.993 

3.434 

Percent  head  to  empty  weight 

1.723 

2.903 

2.719 

1.804 

2.679 

2.899 

Percent  tail  to  empty  weight 

0.087 

0.201 

0.198 

0.105 

0.177 

0.206 

Studies  In  Animal  Nutrition — II 


65 


Table  26. — Distribution  of  Lean,  Fat  and  Bone. 


Steer 

556 

554 

555 

557 

552 

548 

Age 

3 mo. 

3 mo. 

3 mo. 

5 mo. 
17  da. 

5 mo. 
7 da. 

5 mo. 
9 da. 

Group 

1 

2 

3 

1 

2 

3 

Proportion  of  lean,  fat  and  bone  in  empty  animal 


Percent  skeleton 

21.148 

24.090 

23.364 

17.020 

21.443 

22.973 

Percent  lean  flesh 

41.874 

43.747 

44.125 

39.045 

42.579 

43.234 

Percent  fatty  tissue  (excl.  offal  fat) 

4.168 

3.440 

2.334 

13.634 

5.552 

3.114 

Percent  total  fatty  tissue 

5.596 

4.265 

2.994 

17.545 

7.348 

4.097 

Percent  offal  and  kidney  fats  to  total  fatty 
tissue 

33.176 

26.547 

27.820 

32.935 

31.288 

32.047 

Proportion  of  iean,  fat  and  bone  in  carcass 


Percent  skeleton 

25.527 

27.595 

27.343 

19.392 

24.485 

26.535 

Percent  lean  flesh 

66.551 

65.628 

67.372 

59.293 

66.436 

67.631 

Percent  fatty  tissue 

6.535 

4.997 

3.357 

20.384 

6.357 

4.793 

Percent  kidney  fat  to  fatty  tissue  in  carcass. . 

10.611 

9.434 

8.541 

14.159 

9.601 

11.094 

Table  27. — Distribution  of  Lean,  Fat  and  Bone. 


Steer 

547 

550 

558 

541 

538 

540 

Age 

8 mo. 

8 mo. 

8 mo. 

10  mo. 

10  mo. 

11  mo. 

5 da. 

14  da. 

12  da. 

22  da. 

26  da. 

2 da. 

Group 

1 

2 

3 

1 

2 

3 

Proportion  of  lean,  fat  and  bone  in  empty  animal 


Percent  skeleton 

16.096 

19.271 

23.734 

13.301 

17.502 

18.126 

Percent  lean  flesh 

44.646 

42.361 

42.627 

42.917 

43.496 

42.316 

Percent  fatty  tissue  (excl.  offal  fat) 

10.642 

6.916 

3.558 

12.988 

7.139 

6.189 

Percent  total  fattv  tissue 

Percent  offal  and  kidney  fats  to  total  fatty 

12.905 

9.071 

4.479 

16.574 

9.311 

7.864 

tissue 

24.899 

30.639 

26.172 

34.309 

27.534 

27.597 

Proportion  of  lean,  fat  and  bone  in  carcass 


Percent  skeleton 

17.789 

22.864 

27.504 

14.958 

20.418 

21.638 

Percent  lean  flesh 

65.640 

65.534 

65.973 

64.728 

67.616 

67.595 

Percent  fatty  tissue 

15.485 

10.573 

5.159 

19.565 

10.999 

9.636 

Percent  kidney  fat  to  fatty  tissue  in  carcass. . 

9.353 

9.464 

7.820 

16.399 

5.630 

8.341 

Table  28. — Distribution  of  Lean,  Fat  and  Bone. 


Steer 

505 

503 

532 

531 

504 

523 

Age 

10  mo. 

11  mo. 

1 yr.  5 mo. 

1 yr.  6 mo. 

1 yr.  8 mo. 

2 yr.  2 mo. 

18  da. 

11  da. 

20  da. 

12  da. 

26  da. 

6 da. 

Group 

1 

2 

1 

3 

1 

2 

Proportion  of  lean,  fat  and  bone  in  empty  animal 


Percent  skeleton 

13.758 

17.393 

13.557 

17.218 

12.082 

15.161 

Percent  lean  flesh 

41.235 

41.941 

41.765 

43.867 

38.281 

44.821 

Percent  fatty  tissue  (excl.  offal  fat) 

13.662 

8.861 

16.111 

5.158 

18.905 

9.008 

Percent  total  fatty  tissue 

18.321 

11.985 

21.274 

6.668 

24.181 

11.351 

Percent  offal  and  kidney  fats  to  total  fatty 
tissue 

36.875 

33.566 

36.283 

28.314 

31.726 

38.754 

Proportion  of  lean,  fat  and  bone  in  carcass 


Percent  skeleton 

15,978 

20.207 

15.544 

20.686 

18.696 

17.332 

Percent  lean  flesh 

62 . 622 

65.374 

60.392 

70.033 

56.850 

67.631 

Percent  fatty  tissue 

20.576 

13  407 

23.177 

8.130 

28.300 

13.687 

Percent  kidney  fat  to  fatty  tissue  in  carcass. . 

15.716 

10.609 

16.185 

7.553 

12.773 

10.338 

66 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  29. — Distribution  of  Lean,  Fat  and  Bone. 


Steer 

525 

515 

507 

527 

526 

524 

Age 

2 yr.  2 mo. 
8 da. 

2 yr.  9 mo. 
19  da. 

2 yr.  9 mo. 
16  da. 

3 yr.  3 mo. 
15  da. 

3 yr.  4 mo. 

3 yr.  4 mo. 
13  da. 

Group 

3 

1 

2 

1 

2 

3 

Proportion  of  lean,  fat  and  bone  in  empty  animal 


Percent  skeleton 

16.230 

11.555 

15.532 

9.382 

16.138 

19.740 

Percent  lean  flesh 

44.504 

33.109 

44.151 

34.603 

44.777 

46.354 

Percent  fatty  tissue  (excl.  offal  fat) 

6.201 

26.547 

10.631 

31.891 

11.653 

4.242 

Percent  total  fatty  tissue 

Percent  offal  and  kidney  fats  to  total  fatty 

8.069 

30.994 

13.331 

38.063 

14.352 

5.795 

tissue 

29.021 

19.111 

26.085 

22.555 

24.054 

30.913 

Proportion  of  lean,  fat  and  bone  in  carcass 


Percent  skeleton 

19.522 

13.009 

18.004 

9.943 

18.316 

23.425 

Percent  lean  flesh 

69.778 

47.947 

65.918 

46.403 

64.347 

69.762 

Percent  fatty  tissue 

9.625 

38.452 

15.607 

43.137 

16.696 

6.134 

Percent  kidney  fat  to  fatty  tissue  in  carcass . . 

7.894 

5.645 

10.139 

7.605 

6.578 

5.936 

Table  30. — Distribution  of  Lean,  Fat  and  Bone. 


Steer 

513 

502 

509 

501 

512 

500 

Age 

3 yr.  8 mo. 
15  da. 

3 yr.  8 mo. 
19  da. 

3 yr.  8 mo. 
22  da. 

3yr.  11  mo. 

3yr.  11  mo. 
21  da. 

3yr.  11  mo. 
26  da. 

Group 

1 

2 

3 

1 

2 

3 

Proportion  of  lean,  fat  and  bor.e  in  empty  animal 


Percent  skeleton 

9.991 

16.302 

16.570 

9.891 

16.285 

17.509 

Percent  lean  flesh 

34.251 

46.190 

47.013 

31.915 

40 . 983 

45.112 

Percent  fatty  tissue  (excl.  offal  fat) 

30.090 

9.806 

8.710 

35.389 

15.271 

8.307 

Percent  total  fatty  tissue 

37.063 

12.365 

11.245 

40.129 

18.805 

11.480 

Percent  offal  and  kidney  fats  to  total  fatty 
tissue 

23.884 

26.009 

26.121 

17.788 

23.897 

32.832 

Proportion  of  lean,  fat  and  bone  in  carcass 


Percent  skeleton 

11.001 

18.072 

18.747 

10.218 

18.688 

20.361 

Percent  lean  flesh 

46.698 

65.672 

67.804 

42.060 

58.658 

66.597 

Percent  fatty  tissue 

41.405 

14.046 

12.637 

47.340 

22.025 

12.325 

Percent  kidney  fat  to  fatty  tissue  in  carcass. . 

6.283 

6.766 

4.668 

6.794 

6.351 

7.274 

Table  31. — Distribution  of  Lean  Flesh  in  the  Animal. 


Steer 

556 

554 

555 

557 

552 

548 

Age 

3 mo. 

3 mo. 

3 mo. 

5 mo. 

5 mo. 

5 mo. 

17  da. 

7 da. 

9 da. 

Group 

1 

2 

3 

1 

2 

3 

Weight  of  lean  in  animal 

20,546 

17,077 

15,681 

33,734 

21,151 

18,588 

Percent  of  total  lean  in  head 

1.708 

1.956 

2.442 

1.553 

1.882 

2.679 

Percent  of  total  lean  in  shin 

4.088 

3.777 

5.287 

3.436 

4.085 

3.954 

Percent  of  total  lean  in  neck 

1.674 

1.353 

1.830 

1.254 

1.258 

1.544 

Percent  of  total  lean  in  chuck 

26.599 

28.079 

27.409 

26.036 

26.538 

27.077 

Percent  of  total  lean  in  plate 

10.201 

8.637 

9.317 

10.998 

8.922 

8.904 

Percent  of  total  lean  in  rib 

7.841 

8.532 

8.010 

9.444 

9.432 

8.446 

Percent  of  total  lean  in  loin 

16.S84 

17.017 

14.725 

17.362 

16.628 

15.246 

Percent  of  total  lean  in  flank 

2.915 

2.249 

2.889 

3.311 

2.823 

2.394 

Percent  of  total  lean  in  rump 

2.667 

2.653 

2.557 

2.413 

2.605 

2.609 

Percent  of  total  lean  in  round 

23.051 

23.183 

22.722 

22.173 

23.238 

24.769 

Percent  of  total  lean  in  shank 

2.176 

2.342 

2.659 

1.862 

2.340 

2.233 

Percent  of  total  lean  in  tail 

0.195 

0.223 

0.153 

0.157 

0.251 

0.145 

Studies  In  Animal  Nutrition — II 


67 


Table  32. — Distribution  of  Lean  Flesh  in  the  Animal. 


Steer 

547 

550 

558 

541 

538 

540 

Age 

8 mo. 

8 mo. 

8 mo. 

10  mo. 

10  mo. 

11  mo. 

5 da. 

14  da. 

12  da. 

22  da. 

26  da. 

2 da. 

Group 

1 

2 

3 

1 

2 

3 

Weight  of  lean  in  animal 

38.272 

25,652 

19,182 

61,864 

34,560 

29,140 

Percent  of  total  lean  in  head 

3.308 

3.321 

3.618 

1.122 

1.617 

1.527 

Percent  of  total  lean  in  shin 

3.601 

3.863 

4.056 

3.115 

3.464 

4.005 

Percent  of  total  lean  in  neck 

0.792 

1.010 

1.090 

1.578 

1.455 

1.551 

Percent  of  total  lean  in  chuck 

26.390 

26.820 

27.854 

27.271 

27.321 

26.668 

Percent  of  total  lean  in  plate 

9.338 

9.450 

7.226 

10.159 

9.251 

9.602 

Percent  of  total  lean  in  rib 

8.570 

8.124 

8.305 

9.366 

7.517 

8.102 

Percent  of  total  lean  in  loin 

17.736 

17.683 

15.541 

18.925 

18.420 

18.360 

Percent  of  total  lean  in  flank 

3.585 

2.557 

2.409 

2.699 

2.781 

2.255 

Percent  of  total  lean  in  rump 

1.934 

2.460 

2.414 

2.227 

2.257 

2.622 

Percent  of  total  lean  in  round 

22.330 

22.844 

25.190 

21.822 

23.617 

23.089 

Percent  of  total  lean  in  shank 

2.237 

1.696 

2.252 

1.586 

2.182 

2.159 

Percent  of  total  lean  in  tail 

0.180 

0.172 

0.047 

0.129 

0.119 

0.062 

Table  33. — Distribution  of  Lean  Flesh  in  the  Animal. 


Steer 

505 

503 

532 

531 

504 

523 

m 

525 

Age 

10  mo. 

11  mo. 

1 yr.  5 mo. 

1 yr.  6 mo. 

1 yr.  8 mo. 

2 yr.  2 mo. 

2 yr.  2 mo. 

18  da. 

11  da. 

20  da. 

12  da. 

26  da. 

6 da. 

8 da. 

Group 

1 

2 

1 

3 

1 

2 

3 

Weight  of  lean  in  animal 

56,566 

49,580 

95,857 

42,113 

91,081 

75,704 

59,099 

Percent  of  total  lean  in  head. . . 

1.321 

1.743 

1.297 

1.555 

1.458 

1.651 

1.758 

Percent  of  total  lean  in  shin.  . . 

3.566 

3.235 

3.697 

3.804 

3.487 

3.528 

4.097 

Percent  of  total  lean  in  neck. . . 

1.133 

1.105 

0.875 

2.208 

0.927 

1.246 

1.734 

Percent  of  total  lean  in  chuck. . 

28.068 

28.318 

28.167 

27.364 

25.353 

28.695 

26.405 

Percent  of  total  lean  in  plate. . . 

10.361 

8.871 

10.450 

8.513 

10.559 

10.176 

9.562 

Percent  of  total  lean  in  rib 

9.988 

9.008 

9.053 

7.449 

10.159 

7.947 

9.870 

Percent  of  total  lean  in  loin 

17.401 

17.352 

18.849 

16.715 

18.487 

17.063 

15.829 

Percent  of  total  lean  in  flank. . . 

3.380 

2.985 

3.061 

2.044 

4.148 

3.010 

2.883 

Percent  of  total  lean  in  rump . . 

2.298 

2.041 

2.681 

2.377 

3.166 

2.376 

2.689 

Percent  of  total  lean  in  round . . 

20.493 

23.193 

19.855 

25.522 

20.442 

22.390 

23.286 

Percent  of  total  lean  in  shank. . 

1.801 

1.999 

1.850 

2.306 

1.692 

1.749 

1.736 

Percent  of  total  lean  in  tail. . . . 

0.189 

0.151 

0.166 

0.143 

0.122 

0.169 

0.151 

Table  34. — Distribution  of  Lean  Flesh  in  the  Animal. 


Steer 

515 

507 

529 

527 

526 

524 

Age 

2 yr.  9 mo. 
19  da. 

2 yr.  9 mo. 
16  da. 

3 yr.  2 mo. 
21  da. 

3 yr.  3 mo. 
15  da. 

3 yr.  4 mo. 

3 yr.  4 mo. 
13  da. 

Group 

1 

2 

1 

1 

2 

3 

Weight  of  lean  in  animal 

Percent  of  total  lean  in  head 

Percent  of  total  lean  in  shin 

Percent  of  total  lean  in  neck 

Percent  of  total  lean  in  chuck 

Percent  of  total  lean  in  plate 

Percent  of  total  lean  in  rib 

Percent  of  total  lean  in  loin 

Percent  of  total  lean  in  flank 

Percent  of  total  lean  in  rump 

Percent  of  total  lean  in  round 

Percent  of  total  lean  in  shank 

Percent  of  total  lean  in  tail 

111,232 

1.344 

3.436 

0.998 

27.936 

11.857 

8.548 

18.709 

2.950 

3.095 

19.303 

1.684 

0.140 

92,474 

1.281 

3.805 

1.079 

29.534 

10.883 

8.536 

16.072 

2.659 

2.726 

21.250 

2.016 

0.158 

135,989 

1.236 

3.214 

0.536 

26.590 

13.020 

9.508 

18.435 

3.547 

3.272 

18.897 

1.610 

0.133 

95,822 

1.266 

3.866 

1.093 

26.589 

11.208 

9.008 

16.405 

2.339 

2.926 

23.280 

1.849 

0.171 

74,684 

1.595 

3.523 

1.208 

28.576 

8.768 

8.573 

16.201 

2.093 

2.164 

25.249 

1.904 

0.146 

68 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  35. — Distribution  of  Lean  Flesh  in  the  Animal. 


Steer 

513 

502 

509 

501 

512 

500 

3 yr.  8 mo. 

3 yr.  8 mo. 

3 yr.  8 mo. 

3yr.  11  mo. 

3yr.  11  mo 

.3yr.  11  mo 

15  da. 

19  da. 

22  da. 

21  da. 

26  da. 

Group 

1 

2 

3 

1 

2 

3 

Weight  of  lean  in  animal 

132,060 

102,639 

92,018 

130,041 

101,203 

91,990 

Percent  of  total  lean  in  head 

1.397 

1.649 

1.389 

1.540 

1.613 

1.593 

Percent  of  total  lean  in  shin 

3.513 

3.623 

4.096 

3.591 

3.881 

4.565 

Percent  of  total  lean  in  neck 

0.747 

0.920 

1.005 

0.641 

0.790 

0.950 

Percent  of  total  lean  in  chuck 

27.632 

29.227 

28.411 

27.313 

29.911 

28.457 

Percent  of  total  lean  in  plate 

13.834 

9.797 

10.323 

13.913 

11.443 

11.578 

Percent  of  total  lean  in  rib 

9.368 

8.893 

8.890 

8.011 

8.354 

7.393 

Percent  of  total  lean  in  loin 

16.852 

17.101 

16.755 

17.685 

15.840 

16.139 

Percent  of  total  lean  in  flank 

3.208 

2.271 

2.301 

3.494 

1.825 

3.049 

Percent  of  total  lean  in  rump 

2.733 

2.921 

2.859 

2.934 

3.018 

2.686 

Percent  of  total  lean  in  round 

19.227 

21.642 

21.939 

19.275 

21.446 

21.686 

Percent  of  total  lean  in  shank 

1.373 

1.768 

1.856 

1.413 

1.697 

1.683 

Percent  of  total  lean  in  tail 

0.115 

0.187 

0.176 

0.191 

0.181 

0.221 

Table  36. — Distribution  of  Fat  Flesh  in  the  Animal. 


Steer 

556 

554 

555 

557 

552 

548 

Age 

3 mo. 

3 mo. 

3 mo. 

5 mo. 
17  da. 

5 mo. 
7 da. 

5 mo. 
9 da. 

Group 

1 

2 

3 

1 

2 

3 

Weight  of  fat  in  animal 

Percent  of  total  fat  in  head 

Percent  of  total  fat  in  shin 

Percent  of  total  fat  in  neck 

Percent  of  total  fat  in  chuck 

Percent  of  total  fat  in  plate 

Percent  of  total  fat  in  rib  

2,045 

3.227 

3.961 

3.472 

15.795 

7.628 

1.907 

10.269 

20.049 

11.540 

3.619 

16.773 

1.760 

1,343 

5.287 

6.031 

3.797 

13.999 

9.308 

833 

8.643 

8.523 

13.926 

5.042 

11,780 

3.090 

1.647 

1.265 

14.584 

15.017 

6.859 

13.701 

18.981 

8.922 

4.049 

10.747 

0.993 

0.144 

2.758 

5.584 

2.538 

1.958 

14.822 

10.442 

2.067 

9.065 

18.891 

11.965 

4.061 

16.461 

2.139 

1,339 

4.406 

3.809 

4.108 

14.414 

9.783 

1.494 

10.605 

14.339 

12.696 

2.689 

19.417 

2.240 

Percent  of  total  fat  in  kidney  fat 

Percent  of  total  fat  in  loin 

Percent  of  total  fat  in  flank 

Percent  of  total  fat  in  rump 

Percent  of  total  fat  in  round 

Percent  of  total  fat  in  shank 

Percent  of  total  fat  in  tail  . 

8.935 
15.934 

9.978 

5.436 

19.360 

1.936 

7.803 

16.447 

9.364 

4.442 

22.569 

3.241 

Table  37. — Distribution  of  Fat  Flesh  in  the  Animal. 


Steer 

547 

550 

558 

541 

538 

540 

8 mo. 

8 mo. 

8 mo. 

10  mo. 

10  mo. 

11  mo. 

5 da. 

14  da. 

12  da. 

22  da. 

26  da. 

2 da. 

Group 

1 

2 

3 

1 

2 

3 

Weight  of  fat  in  animal 

9,123 

4,188 

1,601 

18,722 

5,672 

4,262 

Percent  of  total  fat  in  head 

4.428 

4.585 

9.744 

1.229 

2.468 

4.036 

Percent  of  total  fat  in  shin 

1.852 

1.552 

3.186 

1.699 

1.693 

1.619 

Percent  of  total  fat  in  neck 

1.261 

0.955 

1.249 

0.977 

2.891 

0.469 

Percent  of  total  fat  in  chuck 

14.250 

17.383 

16.802 

11.804 

18.212 

17.500 

Percent  of  total  fat  in  plate 

13.592 

10.697 

8.495 

13.155 

14.616 

15.157 

Percent  of  total  fat  in  rib 

6.040 

2.865 

1.749 

6.500 

3.755 

3.801 

Percent  of  total  fat  in  kidney  fat 

8.933 

9.026 

7.058 

16.174 

5.483 

8.001 

Percent  of  total  fat  in  loin 

21.769 

25.478 

18.738 

21.600 

20.804 

27.076 

Percent  of  total  fat  in  flank 

10.161 

10.029 

6.996 

10.880 

9.362 

7.297 

Percent  of  total  fat  in  rump 

4.253 

4.776 

3.248 

4.535 

4.725 

5.115 

Percent  of  total  fat  in  round 

11.783 

11.270 

20.112 

10.293 

15.004 

9.503 

Percent  of  total  fat  in  shank 

1.622 

1.337 

2.623 

1.010 

0.846 

0.375 

Percent  of  total  fat  in  tail 

0.055 

0.048 

0.144 

0.141 

0.047 

Studies  In  Animal  Nutrition — II 


69 


Table  38. — Distribution  of  Fat  Flesh  in  the  Animal. 


Steer 

505 

503 

532 

531 

504 

523 

525 

Age 

10  mo. 

11  mo. 

1 yr.  5 mo. 

1 yr.  6 mo. 

1 yr.  8 mo. 

2 yr.  2 mo. 

2 yr.  2 mo. 

18  da. 

11  da. 

20  da. 

12  da. 

26  da. 

6 da. 

8 da. 

Group 

1 

2 

1 

3 

1 

2 

3 

Weight  of  fat  in  animal 

18,742 

10  475 

36.977 

4,952 

44,980 

15,214 

8,234 

Percent  of  total  fat  in  head 

2.172 

4.229 

1.901 

2.827 

0.720 

1.006 

3.085 

Percent  of  total  fat  in  shin 

1.094 

1.403 

1.266 

2.262 

1.670 

1.729 

2.417 

Percent  of  total  fat  in  neck 

0.720 

1.728 

0.749 

2.383 

0.518 

0.381 

1.421 

Percent  of  total  fat  in  chuck. . . . 

13.440 

11.437 

13.484 

17.750 

12.750 

16.012 

14.149 

Percent  of  total  fat  in  plate 

13.798 

14.081 

15.299 

12.298 

15.480 

13.330 

14.258 

Percent  of  total  fat  in  rib 

7.043 

4.010 

8.375 

3.413 

7.526 

5.002 

4.032 

Percent  of  total  fat  in  kidney  fat 

15.351 

10.148 

15.867 

7.330 

12.672 

10.221 

7.639 

Percent  of  total  fat  in  loin 

20.163 

27.427 

20.221 

22.859 

20.387 

20.954 

22.820 

Percent  of  total  fat  in  flank 

9.273 

7.561 

10.033 

7.512 

9.931 

9.734 

10.347 

Percent  of  total  fat  in  rump .... 

4.877 

5.642 

3.946 

5.089 

5.645 

5.981 

5.927 

Percent  of  total  fat  in  round .... 

11.050 

10.587 

8.200 

15.044 

10.914 

14.973 

11.914 

Percent  of  total  fat  in  shank 

0.864 

1.632 

0.592 

1.111 

1.716 

0.552 

1.846 

Percent  of  total  fat  in  tail 

0.155 

0.115 

0.068 

0.121 

0.071 

0.125 

0.146 

Table  39. — Distribution  of  Fat  Flesh  in  the  Animal. 


Steer 

515 

507 

529 

527 

526 

524 

Age 

2 yr.  9 mo. 
16  da. 

3 yr.  2 mo. 
21  da. 

3 yr.  3 mo. 
15  da. 

3 yr.  4 mo. 

3 yr.  4 mo. 
13  da. 

19  da. 

Group 

1 

2 

1 

1 

2 

3 

Weight  of  fat  in  animal 

Percent  of  total  fat  in  head 

Percent  of  total  fat  in  shin 

Percent  of  total  fat  in  neck 

Percent  of  total  fat  in  chuck 

Percent  of  total  fat  in  plate 

Percent  of  total  fat  in  rib 

Percent  of  total  fat  in  kidney  fat 

Percent  of  total  fat  in  loin 

Percent  of  total  fat  in  flank 

Percent  of  total  fat  in  rump 

Percent  of  total  fat  in  round 

Percent  of  total  fat  in  shank 

Percent  of  total  fat  in  tail 

89,188 

1.393 

1.724 

0.748 

12.576 

20.570 

9.129 

5.562 

21.485 

9.814 

5.003 

10.684 

1.239 

0.074 

22,267 

3.036 

1.159 

0.804 

12.471 

15.224 

5.461 

9.826 

22.877 

9.665 

6.099 

12.076 

1.298 

0.054 

125,332 

0.497 

1.022 

0.481 

14.943 

20.466 

9.685 

7.566 

21.034 

9.024 

6.032 

8.564 

0.668 

0.019 

24,937 

1.648 

0.995 

1.335 

15.062 

16.482 

7.459 

6.464 

23.327 

10.711 

5.987 

10.057 

0.393 

0.080 

6,834 
5.414 
2.766 
0 556 
13.330 
14.969 
2.473 
5.604 
17.881 
11.121 
5.838 
18.481 
1.390 
0.176 

Table  40. — Distribution  of  Fat  Flesh  in  the  Animal. 


Steer 

513 

502 

509 

501 

512 

500 

Age 

3 yr.  8 mo. 
15  da. 

3 yr.  8 mo. 
19  da. 

3 yr.  8 mo. 
22  da. 

3yr.  11  mo. 

3yr.  11  mo. 
21  da. 

3 yr.  1 1 mo 
26  da. 

Group 

1 

2 

3 

1 

2 

3 

Weight  of  fat  in  animal 

116,017 

21,789 

17,048 

144,196 

37,709 

16,940 

Percent  of  total  fat  in  head 

0.558 

1.005 

0.774 

0.210 

0.936 

1.122 

Percent  of  total  fat  in  shin 

0.896 

1 629 

1.396 

0.848 

1.223 

1.865 

Percent  of  total  fat  in  neck 

0.531 

0.863 

1.003 

0.387 

0.464 

1.169 

Percent  of  total  fat  in  chuck 

14.977 

19.469 

16.712 

12.890 

14.612 

13.040 

Percent  of  total  fat  in  plate 

20.796 

15.659 

16.987 

21.006 

19.820 

18.017 

Percent  of  total  fat  in  rib 

10.174 

7.660 

5.801 

9.821 

7.157 

5.325 

Percent  of  total  fat  in  kidney  fat 

6.245 

6.691 

4.622 

6 777 

6.285 

7.178 

Percent  of  total  fat  in  loin 

21.518 

20.983 

22.202 

24.743 

20.298 

20.159 

Percent  of  total  fat  in  flank 

9.700 

9.170 

8.112 

9.810 

9.470 

10  018 

Percent  of  total  fat  in  rump 

5.113 

4.828 

6.183 

5.060 

5.802 

6.246 

Percent  of  total  fat  in  round 

8.235 

10.602 

14.975 

7.727 

13.180 

14.569 

Percent  of  total  fat  in  shank 

1.214 

1.345 

1.032 

0.680 

0.655 

1.092 

Percent  of  total  fat  in  tail 

0.044 

0.096 

0.199 

0.041 

0.098 

0.201 

70 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  41. — Distribution  of  Skeleton  in  the  Animal. 


Steer 

556 

554 

555 

557 

552 

548 

Age 

3 mo. 

3 mo. 

3 mo. 

5 mo. 
17  da. 

5 mo. 
7 da. 

5 mo. 
9 da. 

Group 

1 

2 

3 

1 

2 

3 

Wt.  of  skeleton  in  animal 

10,377 

9,404 

8,304 

14,705 

10,652 

9,877 

Percent  of  total  skeleton  in  head 

12.576 

11.857 

11.911 

13.499 

14.327 

15.136 

Percent  of  total  skeleton  in  shin 

8.201 

7.784 

8.755 

7.637 

7.276 

8.019 

Percent  of  total  skeleton  in  neck 

0.964 

1.414 

2.589 

1.401 

1.070 

1.934 

Percent  of  total  skeleton  in  chuck 

17.029 

18.429 

17.438 

17.769 

17.275 

16.442 

Percent  of  total  skeleton  in  plate 

7.450 

6.678 

7.286 

8.439 

7.041 

7.502 

Percent  of  total  skeleton  in  rib 

7.931 

7.199 

7.563 

8.745 

9.783 

8.170 

Percent  of  total  skeleton  in  loin 

10.929 

9.932 

7.611 

8.426 

10.290 

8.363 

Percent  of  total  skeleton  in  flank 

0.154 

0.117 

0.157 

0.054 

0.075 

0.162 

Percent  of  total  skeleton  in  rump 

2.930 

3.892 

3.047 

3.264 

3.004 

2.885 

Percent  of  total  skeleton  in  round 

9.512 

9.050 

9.948 

9.133 

7.633 

9.608 

Percent  of  total  skeleton  in  shank 

9.406 

10.198 

10.261 

8.875 

8.177 

8.667 

Percent  of  total  skeleton  in  tail 

0.443 

0.670 

0.446 

0.660 

0.620 

0.476 

Percent  of  total  skeleton  in  feet 

12.475 

12.777 

12.988 

12.098 

13.430 

12.635 

Table  42. — Distribution  of  Skeleton  in  the  Animal. 


Steer 

547 

550 

558 

541 

538 

540 

Age 

8 mo. 
5 da. 

8 mo. 
14  da. 

8 mo. 
12  da. 

10  mo. 
22  da. 

10  mo. 
26  da. 

11  mo. 
2 da. 

Group 

1 

2 

3 

1 

2 

3 

Weight  of  skeleton  in  animal 

13,798 

11,670 

10,680 

19,173 

13,906 

12,482 

Percent  of  total  skeleton  in  head 

11.893 

13.539 

14.869 

14.082 

14.389 

14.749 

Percent  of  total  skeleton  in  shin 

6.726 

7.506 

8.090 

7.683 

7.939 

7.819 

Percent  of  total  skeleton  in  neck 

0.790 

1.260 

0.964 

1.763 

1.438 

1.706 

Percent  of  total  skeleton  in  chuck 

17.763 

19.974 

18.024 

17.264 

18.021 

17.906 

Percent  of  total  skeleton  in  plate 

7.023 

6.530 

5.075 

7.380 

7.637 

7.395 

Percent  of  total  skeleton  in  rib 

7.871 

7.455 

7.097 

8.246 

7.745 

7.307 

Percent  of  total  skeleton  in  loin 

11.349 

10.557 

10.009 

10.212 

9.694 

10.215 

Percent  of  total  skeleton  in  flank 

0.072 

0.069 

0.037 

0.089 

0.072 

0.064 

Percent  of  total  skeleton  in  rump 

2.754 

2.999 

3.174 

2.754 

2.632 

2.492 

Percent  of  total  skeleton  in  round 

9.016 

8.997 

9.803 

8.475 

9.190 

9.414 

Percent  of  total  skeleton  in  shank 

9.182 

8.663 

9.860 

9.764 

9.377 

9.229 

Percent  of  total  skeleton  in  tail 

0.522 

0.403 

0.375 

0.537 

0.489 

0.553 

Percent  of  total  skeleton  in  feet 

15.038 

12.048 

12.622 

11.751 

11.384 

11.152 

Table  43. — Distribution  of  Skeleton  in  the  Animal. 


Steer 

505 

503 

532 

531 

504 

523 

525 

10  mo. 

11  mo. 

1 yr.  5 mo. 

1 yr.  6 mo. 

1 yr.  8 mo. 

2 yr.  2 mo. 

2 yr.  2 mo. 

Age 

18  da. 

11  da. 

20  da. 

12  da. 

26  da. 

6 da. 

8 da. 

Group 

1 

2 

1 

3 

1 

2 

3 

Weight  of  skeleton  in  animal 

18,873 

20,561 

31,116 

16,539 

28,747 

25,608 

21,552 

Percent  of  total  skeleton  in  head  . 

12.876 

14.377 

11.014 

14.198 

13.988 

14.128 

13.943 

Percent  of  total  skeleton  in  shin. . 

7.837 

7.626 

8.687 

8.427 

7.528 

7.791 

7.800 

Percent  of  total  skeleton  in  neck. . 

1.399 

2.091 

1.189 

1.555 

1.579 

1.476 

1.847 

Percent  of  total  skeleton  in  chuck. 

18.529 

16.458 

19.723 

16.891 

17.859 

17.631 

17.757 

Percent  of  total  skeleton  in  plate. 

7.831 

8.151 

8.642 

7.550 

7.823 

7.474 

7.865 

Percent  of  total  skeleton  in  rib. . . 

8.457 

8.224 

8.391 

6.897 

8.857 

7.505 

9.020 

Percent  of  total  skeleton  in  loin . . 

10.263 

9.323 

10.037 

8.572 

10.175 

10.391 

9.113 

Percent  of  total  skeleton  In  flank . 

0.106 

0.204 

0.161 

0.151 

0.129 

0.105 

0.148 

Percent  of  total  skeleton  in  rump. 

2.930 

2.553 

3.667 

3.206 

4.223 

3.456 

3.577 

Percent  of  total  skeletonin  round . 

8.997 

9.377 

9.037 

11.010 

8.363 

9.040 

9.387 

Percent  of  total  skeleton  in  shank. 

8.970 

9.114 

8.597 

9.716 

8.589 

9.513 

8.473 

Percent  of  total  skeleton  in  tail . . . 

0.636 

0.462 

0.427 

0.448 

0.550 

0.617 

0.520 

Percent  of  total  skeleton  in  feet. . 

11.169 

12.037 

10.429 

11.379 

10.338 

10.872 

10.551 

Studies  In  Animal  Nutrition — II 


71 


Table  44. — Distribution  of  Skeleton  in  the  Animal  . 


515 

507 

529 

527 

526 

524 

2 yr.  9 mo. 
19  da. 

2 yr.  9 mo. 
16  da. 

3 yr.  2 mo. 
21  da. 

3 yr.  3 mo. 
15  da. 

3 yr.  4 mo. 

3 yr.  4 mo 
13  da. 

Group 

1 

2 

1 

1 

2 

3 

Weight  of  skeleton  in  animal 

Percent  of  total  skeleton  in  head 

Percent  of  total  skeleton  in  shin 

Percent  of  total  skeleton  in  neck 

Percent  of  total  skeleton  in  chuck 

Percent  of  total  skeleton  in  plate 

Percent  of  total  skeleton  in  rib 

Percent  of  total  skeleton  in  loin 

Percent  of  total  skeleton  in  flank 

Percent  of  total  skeleton  in  rump 

Percent  of  total  skeleton  in  round 

Percent  of  total  skeleton  in  shank 

Percent  of  total  skeleton  in  tail 

Percent  of  total  skeleton  in  feet 

38,821 

12.725 

7.751 

1.309 

17.403 

8.047 

8.325 

10.026 

0.155 

5.247 

8.171 

10.152 

0.456 

10.234 

32,531 

13.249 

7.716 

1.282 

19.010 

9.425 

7.762 

10.000 

0.224 

3.898 

9.013 

8.192 

0.544 

9.686 

36,872 

11.491 

8.597 

1.378 

18.787 

8.188 

8.877 

9.682 

0.060 

4.421 

8.741 

9.216 

0.502 

10.092 

34,535 

12.790 

8.166 

1.231 

18.810 

9.315 

8.238 

9.909 

0.223 

4.109 

9.196 

8.646 

0.481 

8.887 

31,805 

12.621 

7.782 

1.654 

18.620 

9.102 

8.348 

10.354 

0.214 

3.811 

9.241 

8.351 

0.456 

9.448 

Table  45. — Distribution  of  Skeleton  in  the  Animal. 


Steer 

513 

502 

509 

501 

512 

500 

Age 

3 yr.  8 mo. 
15  da. 

3 yr.  8 mo. 
19  da. 

3 yr.  8 mo. 
22  da. 

3yr.  11  mo. 

3yr.  11  mo. 
21  da. 

3yr.  11  mo. 
26  da. 

Group 

1 

2 

3 

1 

2 

3 

Weight  of  skeleton  in  animal 

38,521 

36,226 

32,433 

40,301 

40,214 

35,704 

Percent  of  total  skeleton  in  head 

10.799 

13.220 

12.715 

12.980 

12.018 

12.539 

Percent  of  total  skeleton  in  shin 

7.944 

8.199 

7.779 

7.655 

7.552 

7.856 

Percent  of  total  skeleton  in  neck 

0.927 

1.275 

1.656 

1.109 

1.542 

1.650 

Percent  of  total  skeleton  in  chuck 

19.226 

19.555 

19.437 

18.466 

19.018 

18.586 

Percent  of  total  skeleton  in  plate 

9.405 

7.489 

7.745 

8.652 

9.517 

8.666 

Percent  of  total  skeleton  in  rib 

9.211 

8.171 

7.687 

7.925 

8.626 

7.271 

Percent  of  total  skeleton  in  loin 

11.080 

10.857 

10.594 

10.687 

10.877 

10.884 

Percent  of  total  skeleton  in  flank 

0.249 

0.226 

0.154 

0.117 

0.167 

0.227 

Percent  of  total  skeleton  in  rump 

4.496 

3.260 

4.409 

4.568 

4.058 

4.184 

Percent  ot  total  skeleton  in  round 

9.143 

9.253 

9.281 

9.012 

9.730 

9.968 

Percent  of  total  skeleton  in  shank 

7.863 

8.251 

8.476 

8.843 

7.654 

8.052 

Percent  of  total  skeleton  in  tail 

0.387 

0.610 

0.595 

0.377 

0.517 

0.541 

Percent  of  total  skeleton  in  feet 

9.862 

9.637 

9.472 

9.608 

8.723 

9.576 

72 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  46. — Proportion  of  Lean,  Fat,  and  Bone  in  Cuts  of  the  Carcass. 


Steer 

556 

554 

555 

557 

552 

548 

3 mo. 

3 mo. 

3 mo. 

5 mo. 

5 mo. 

5 mo. 

17  da. 

7 da. 

9 da. 

Group 

j 

2 

3 

1 

2 

3 

47.592 

43.937 

50.641 

47.018 

50.824 

46.168 

4.589 

5.518 

4.337 

7.870 

4.118 

3.204 

48.215 

49.864 

44.411 

45.558 

45.588 

49.749 

66.538 

55.797 

57.171 

55.007 

62.150 

54.459 

13.733 

12.319 

19.376 

12.617 

10.436 

Percent  of  bone  in  neck 

19.342 

32.126 

42.829 

26.788 

26.636 

36.243 

71.908 

71 . 100 

73.307 

67.107 

71.494 

73.303 

Percent  of  fat  in  chuck 

4.250 

2.788 

1.979 

13.127 

5.210 

2.811 

Percent  of  bone  in  chuck 

23.250 

25.697 

24.697 

19.965 

23.437 

23.653 

Percent  of  lean  in  plate 

68.766 

65.702 

69.340 

55.053 

64 . 184 

65.545 

Percent  of  fat  in  plate 

5.118. 

5.568 

1.993 

26.250 

9.796 

5.188 

Percent  of  bone  in  plate 

25.361 

27.973 

28.714 

18.415 

25.510 

29.347 

Percent  of  lean  in  rib 

64.829 

67.957 

65.587 

60.387 

63.942 

65.091 

Percent  of  fat  in  rib 

1.569 

15.315 

1.827 

0.829 

Percent  of  bone  in  rib 

33.119 

31.576 

32.794 

24.375 

33.397 

33.458 

Percent  of  lean  in  loin 

68 . 980 

71.086 

74.436 

62.528 

68.331 

73.344 

Percent  of  fat  in  loin 

8.153 

5.235 

4.417 

23.871 

10.122 

4.969 

Percent  of  bone  in  loin 

22.549 

22.847 

20.374 

13.227 

21.294 

21.377 

Percent  of  lean  in  flank 

69.009 

72.045 

82.514 

51.498 

63.850 

70.635 

Percent  of  fat  in  flank 

27.189 

25.141 

14.208 

48.456 

35.294 

26.984 

Percent  of  bone  in  flank 

1.843 

2.064 

2.368 

0.369 

0.856 

2.540 

Percent  of  lean  in  rump 

59.179 

50.842 

57.041 

45.756 

55.826 

59.803 

Percent  of  fat  in  rump 

7.997 

8.193 

5.263 

28.813 

11.348 

4.439 

Percent  of  bone  in  rump 

32.829 

41.077 

35.989 

26.981 

32.421 

35  142 

Percent  of  lean  in  round 

77.563 

78.179 

77.457 

74.133 

78.892 

79.229 

Percent  of  fat  in  round 

5.617 

5.134 

4.087 

12.547 

7.287 

4.474 

Percent  of  bone  in  round 

16.164 

16.805 

17.957 

13.310 

13.050 

16.331 

Percent  of  lean  in  shank 

30.244 

28.674 

31.856 

30.426 

34.640 

31.801 

Percent  of  fat  in  shank 

2.436 

1.804 

2.063 

5.669 

4.129 

2.299 

Percent  of  bone  in  shank 

66.035 

68.746 

65.088 

63.227 

60.952 

65.594 

Studies  In  Animal  Nutrition — II 


73 


Table  47. — Proportion  of  Lean,  Fat,  and  Bone  in  Cuts  of  the  Carcass. 


547 

550 

558 

541 

538 

540 

8 mo. 

8 mo. 

8 mo. 

10  mo. 

10  mo. 

11  mo. 

5 da. 

14  da. 

12  da. 

22  da. 

26  da. 

2 da 

1 

2 

3 

j 

2 

3 

55.408 

50.794 

45.630 

51.913 

49.875 

52.379 

6.795 

3.332 

2.991 

8.567 

4.000 

3.097 

37.314 

44.900 

50.674 

39.682 

46.000 

43.806 

57.714 

57.428 

60.933 

65.067 

58.762 

65.602 

21.905 

8.869 

5.831 

12.200 

19.159 

2.903 

20.762 

32.594 

30.029 

22.533 

23.364 

30.914 

72.651 

69.062 

70.703 

74.054 

72.430 

72.201 

9.351 

7.308 

3.560 

9.701 

7.924 

6.931 

Percent  of  bone  in  chuck 

17.631 

23.399 

25.473 

14.529 

19.224 

20.766 

Percent  of  lean  in  plate 

61.578 

64.726 

67.445 

61.830 

62.441 

63.969 

Percent  of  fat  in  plate 

21.365 

11.963 

6.618 

24.230 

16.191 

14.769 

Percent  of  bone  in  plate 

16.695 

20.347 

26.375 

13.920 

20.742 

21.102 

Percent  of  lean  in  rib 

66.504 

67.553 

66.933 

67.138 

66.855 

69.075 

Percent  of  fat  in  rib 

11.172 

3.890 

1.176 

14.102 

5.481 

4.740 

Percent  of  bone  in  rib 

22.019 

28.201 

31.849 

18.320 

27.715 

26.682 

68.563 

Percent  of  lean  in  loin 

65.439 

66.316 

68.387 

65.654 

71.073 

Percent  of  fat  in  loin 

19.146 

15.599 

6.882 

22.677 

13.174 

14 . 789 

Percent  of  bone  in  loin 

15.097 

18.012 

24.524 

10.980 

15.050 

16.340 

Percent  of  lean  in  flank 

59.215 

59.582 

80.348 

44.832 

67.015 

67.523 

Percent  of  fat  in  flank 

40.009 

38.147 

19.478 

54.685 

37.029 

31.963 

Percent  of  bone  in  flank 

0.432 

0.727 

0.696 

0.456 

0.697 

0.822 

Percent  of  lean  in  rump 

48.334 

53.475 

53.464 

49.837 

54.318 

58 . 769 

Percent  of  fat  in  rump 

25.343 

16.949 

6.005 

30.705 

18.663 

16.769 

Percent  of  bone  in  rump 

24.820 

29.661 

39.145 

19.096 

25.487 

23 . 923 

Percent  of  lean  in  round 

78.613 

79.286 

77.647 

79.109 

79.289 

81.011 

Percent  of  fat  in  round 

9.889 

6.386 

5.174 

11.292 

8.267 

4.877 

Percent  of  bone  in  round 

11.443 

14.206 

16.825 

9.522 

12.415 

14.148 

Percent  of  lean  in  shank 

37.462 

28.961 

28  107 

32.249 

35.802 

34.751 

Percent  of  fat  in  6hank 

6.477 

3.728 

2.733 

6.213 

2.279 

0.884 

Percent  of  bone  in  shank 

55.449 

67.310 

68.510 

61.538 

61.918 

63.646 

74 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  48. — Proportion  of  Lean,  Fat,  and  Bone  in  Cuts  of  the  Carcass. 


Steer 

505 

503 

532 

531 

504 

523 

525 

Age 

10  mo. 

11  mo. 

1 yr.  5 mo. 

1 yr.  6 mo. 

1 yr.  8 mo. 

2 yr.  2 mo. 

2 yr.  2 mo. 

i§ 

18  da. 

11  da' 

20  da. 

12  da. 

26  da. 

6 da. 

8 da. 

Group 

1 

2 

1 

3 

1 

2 

3 

Percent  of  lean  in  shin 

54.469 

48.168 

52.566 

51.561 

51.870 

54.058 

55.938 

Percent  of  fat  in  shin 

5.536 

4.414 

6.942 

3.605 

12.265 

5.323 

4.598 

Percent  of  bone  in  shin 

39.941 

47.087 

40.092 

44.834 

35.342 

40.376 

38.840 

Percent  of  lean  in  neck 

61.398 

47.282 

56.843 

71.981 

54.948 

67.693 

66.044 

Percent  of  fat  in  neck 

12.931 

15.617 

18.767 

9.133 

15.169 

4.170 

7.539 

Percent  of  bone  in  neck 

25.287 

37.101 

25.068 

19.892 

29.557 

27.175 

25.644 

Percent  of  lean  in  chuck 

72.310 

74.598 

70.518 

75.305 

66.653 

75.017 

75.657 

Percent  of  fat  in  chuck 

11.472 

6.365 

13.022 

5.744 

16.554 

8.430 

5.648 

Percent  of  bone  in  chuck 

15.927 

17.980 

16.029 

18.245 

14.819 

15.624 

18.554 

Percent  of  lean  in  plate 

58.946 

58.021 

54.207 

65.384 

50.870 

64.227 

66.047 

Percent  of  fat  in  plate 

26.008 

19.459 

30.613 

11.107 

36.832 

16.907 

13.721 

Percent  of  bone  in  plate 

14.865 

22.111 

14.552 

22.761 

11.896 

15.957 

19.811 

Percent  of  lean  in  rib 

65.644 

67.411 

59.919 

69.975 

60.532 

68.731 

71.711 

Percent  of  fat  in  rib 

15.336 

6.340 

21.384 

3.770 

22.144 

8.694 

4.082 

Percent  of  bone  in  rib 

18.543 

25.525 

18.028 

25.429 

16.656 

21.958 

23.900 

Percent  of  lean  in  loin 

63.007 

63.844 

66.024 

73.011 

57.821 

68.547 

70.652 

Percent  of  fat  in  loin 

24.190 

21.321 

27.322 

11.742 

31.489 

16.918 

14.191 

Percent  of  bone  in  loin 

12.399 

14.266 

11.412 

14.698 

10.044 

14.121 

14.833 

Percent  of  lean  in  flank 

52.169 

63.519 

43.843 

68.715 

45.628 

60.053 

65.589 

Percent  of  fat  in  flank 

47.422 

33.991 

55.439 

29.689 

53.949 

39.025 

32.794 

Percent  of  bone  in  flank 

0.546 

1.803 

0.747 

1.995 

0.447 

0.711 

1.232 

Percent  of  lean  in  rump 

46.662 

47.158 

49.518 

56.141 

43.493 

49.600 

55.250 

Percent  of  fat  in  rump 

32.807 

27.540 

28.112 

14.133 

38.290 

25.090 

16.968 

Percent  of  bone  in  rump 

19.849 

24.464 

21.985 

29.725 

18.308 

24.400 

26.808 

Percent  of  lean  in  round 

75.371 

78.760 

76.110 

80.515 

71.625 

78.425 

81.708 

Percent  of  fat  in  round 

13.466 

7.596 

12.125 

5.581 

18.884 

10.540 

5.824 

Percent  of  bone  in  round 

11.040 

13.205 

11.245 

13.634 

9.248 

10.711 

12.011 

Percent  of  lean  in  shank 

35.321 

32.567 

37.868 

36.559 

31.991 

34.177 

33.929 

Percent  of  fat  in  shank 

5.615 

5.619 

4.677 

2.071 

16.027 

2.168 

5.026 

Percent  of  bone  in  shank 

58.683 

61.584 

57.134 

60.467 

51.256 

62.881 

60.384 

Studies  In  Animal  Nutrition — II 


75 


Table  49. — Proportion  of  Lean,  Fat,  and  Bone  in  Cuts  of  the  Carcass. 


Steer 

515 

507 

529 

527 

526 

521 

Age 

2 yr.  9 mo. 

2 yr.  9 mo. 

3 yr.  2 mo. 

3 yr.  3 mo. 

3 yr.  4 mo. 

3 yr.  4 mo 

19  da. 

16  da. 

21  da. 

15  da. 

13  da. 

Group 

1 

2 

1 

1 

2 

3 

45.663 

55 . 742 

49.535 

54.559 

49.334 

18.375 

4.087 

14.517 

3.653 

3.544 

35.950 

39.759 

35.925 

41.538 

46.409 

48.514 

61 . 949 

39.792 

57.781 

61.236 

29.152 

11.111 

32.915 

18.377 

2.580 

22.203 

25.885 

27.729 

23.455 

35.709 

63.029 

74.907 

58.273 

70.957 

75.584 

Percent  of  fat  in  chuck 

22.750 

7.617 

30.206 

10.461 

3.510 

Percent  of  bone  in  chuck 

13.704 

16.961 

11.172 

18.092 

20.973 

Percent  of  lean  in  plate 

37.818 

60.616 

37.983 

59.327 

62.445 

Percent  of  fat  in  plate 

52.605 

20.418 

55.024 

22.703 

9.756 

Percent  ot  bone  in  plate 

8.958 

18.467 

6.476 

17.771 

27.608 

Percent  of  lean  in  rib. 

45.259 

67.847 

49.110 

45.527 

64.558 

69.477 

Percent  of  fat  in  rib 

38.757 

10.451 

36.531 

42.742 

13.911 

1.834 

Percent  of  bone  in  rib 

15.385 

21.702 

14.064 

11.524 

21.277 

28.809 

Percent  of  lean  in  loin 

47.464 

63.583 

45.427 

62.855 

72 . 646 

Percent  of  fat  in  loin 

43.705 

21.793 

47.768 

23.259 

7.337 

Percent  of  bone  in  loin 

8.877 

13.917 

6.469 

13.683 

19.771 

Percent  of  lean  in  fla  nk 

27.089 

52.353 

29.832 

45.429 

64.936 

Percent  of  fat  in  flank 

72.267 

45.816 

69.957 

54.146 

31.575 

Percent  of  bone  in  fl  \nk 

0.495 

1.554 

0.136 

1.561 

2.825 

Percent  of  lean  in  rump 

34.413 

49.047 

32.432 

48.782 

50.000 

Percent  of  fat  in  rump 

44.598 

26.206 

55.098 

25.974 

12.345 

Percent  of  bone  in  rump 

20.360 

24.669 

11.880 

24 . 687 

37.500 

Percent  of  lean  in  round 

62.832 

77.752 

64.656 

79.379 

81.941 

Percent  of  fat  in  round 

27.885 

10.639 

27.004 

8.925 

5.488 

Percent  of  bone  in  round 

9.282 

11.601 

8.109 

11.302 

12.771 

Percent  of  lean  in  shank  

27.059 

38.314 

34.038 

36.423 

33.841 

Percent  of  fat  in  shank 

15.964 

5.940 

13.009 

2.014 

2.261 

Percent  of  bone  in  shank 

56.934 

54.779 

52.813 

61.377 

63.208 

76 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  54 


Table  50. — Proportion  of  Lean,  Fat,  and  Bone  in  Cuts  of  the  Carcass. 


513 

502 

509 

501 

512 

500 

3 yr.  8 mo. 
19  da. 

3 yr.  8 mo. 
22  da. 

3yr.  11  mo. 

3yr.  11  mo. 
21  da. 

3yr.  11  mo 
26  da. 

15  da. 

j 

2 

3 

1 

2 

3 

Percent  of  lean  in  shin » 

53.029 

11.900 

52.887 

5.048 

57.780 

3.649 

51.785 

13.562 

52.817 

6.199 

57.090 

4.297 

34.979 

42.235 

38.679 

34.154 

40.836 

38.138 

50.772 

59.557 

56.575 

45.247 

50.000 

52.651 

31 . 687 

11.861 

10.459 

30.310 

10.938 

11.928 

18.364 

29.148 

32.844 

24.280 

38.750 

35.482 

59.413 

72.279 

73.539 

57.534 

69.481 

74.371 

Percent  of  fat  in  chuck 

28.291 

10.221 

8.014 

30.106 

12.647 

6.276 

Percent  of  bone  in  chuck 

12.058 

17.069 

17.735 

12.055 

17.555 

18.853 

Percent  of  lean  in  plate 

39.560 

61.906 

63.653 

34.810 

50.339 

63.406 

Percent  of  fat  in  plate 

52.244 

21.005 

19.406 

58.276 

32.487 

18.169 

Percent  of  bone  in  plate 

7.845 

16.702 

16.833 

6.709 

16.635 

18.419 

Percent  of  lean  in  rib 

44.456 

66.044 

69.765 

37.494 

58.195 

65.894 

Percent  of  fat  in  rib 

42.415 

12.076 

8.435 

50.970 

18.579 

8.7-39 

Percent  of  bone  in  rib 

12.749 

21.417 

21.262 

11.496 

23.880 

25.153 

Percent  of  lean  in  loin 

42.388 

67.110 

67.724 

36.472 

57.149 

66.998 

Percent  of  fat  in  loin 

47.548 

17.481 

16.626 

56.583 

27.286 

15.411 

Percent  of  bone  in  loin 

8.129 

15.038 

15.093 

6.830 

15.593 

17.537 

Percent  of  lean  in  flank 

27.230 

53.098 

59.500 

24.211 

33.570 

61.164 

Percent  of  fat  in  flank 

72.326 

45.513 

38.870 

75.373 

64.904 

37.004 

Percent  of  bone  in  flank 

0.617 

1.868 

1.405 

0.250 

1.218 

1.766 

Percent  of  lean  in  rump 

31.972 

57.181 

51.167 

29.313 

44.454 

49.018 

Percent  of  fat  in  rump 

52.551 

20.065 

20.498 

56.053 

31.849 

20.988 

Percent  of  bone  in  rump 

15.344 

22.525 

27.810 

14.142 

23.755 

29.637 

Percent  of  lean  in  round 

65.686 

79.267 

78.060 

62.710 

70.640 

76.512 

Percent  of  fat  in  round 

24.716 

8.243 

9.872 

27.876 

16.176 

9.466 

Percent  of  bone  in  round 

9.111 

11.962 

11.639 

9.087 

12.736 

13.650 

Percent  of  lean  in  shank 

29.031 

35.749 

36.818 

28.789 

33.859 

33.240 

Percent  of  fat  in  shank 

22.546 

5.771 

3.794 

15.358 

4.871 

3.973 

Percent  of  bone  in  shank 

48.503 

58.873 

59.258 

55.853 

60.698 

61.735 

UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 

AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  55 


Studies  In  Animal  Nutrition 

III.  Changes  in  Chemical  Composition 
on  Different  Planes  of  Nutrition 


COLUMBIA,  MISSOURI 
OCTOBER,  1922 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY  P.  E.  BURTON  H.  J.  BLANTON 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 

J.  C.  JONES,  PH.  D.,  LL.  D.,  PRESIDENT  OF  THE  UNIVERSITY 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 


STATION  STAFF 

OCTOBER,  1922 


AGRICULTURAL  CHEMISTRY 


RURAL  LIFE 
O.  R.  Johnson,  A.  M. 


C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  Ph.  D. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  Sieveking,  B.  S.  in  Agr. 

AGRICULTURAL  ENGINEERING 
J.  C.  Wooley,  B.  S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

DAIRY  HUSBANDRY 
A.  C.  Ragsdale,  B.  S.  in  Agr. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Nelson,  B.  S.  in  Agr. 

W.  P.  Hays 


ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride,  B.  S.  in  Agr. 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  Ph.  D. 

O.  W.  Letson,  B.  S.  in  Agr. 
Miss  Regina  Schulte* 


S.  D.  Gromer,  A.  M. 

E.  L.  Morgan,  A.M. 

Ben  H.  Frame,  B.  S.  in  Agr. 
Owen  Howells,  B.  S.  in  Agr. 


HORTICULTURE 
T.  J.  Talbert,  A.  M. 

H.  D.  Hooker,  Tr.,  Ph.  D. 

J.  T.  Rosa,  Jr.,  Ph.  D. 

H.  G.  Swartwout.  B.  S.  in  Agr. 

J.  T.  Quinn,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY 
H.  L.  Kempster,  B.  S. 

Earl  W.  Henderson,  B.S. 

SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M 
W.  A.  Albrecht,  Ph.  D. 

F.  L.  Duley,  A.M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr. 

Richard  Bradfield,  Ph.  D. 

VETERINARY  SCIENCE 
J.  W.  Connaway,  D.  V.  S.,  M.  D. 

L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 

OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Secretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Jane  Frodsiiam,  Librarian. 

E.  E.  Brown,  Business  Manager. 


In  service  of  U.  S.  Department  of  Agriculture. 


STUDIES  IN  ANIMAL  NUTRITION 

III.  Changes  in  Chemical  Composition  on  Different 
Planes  of  Nutrition. 

C.  Robert  Moulton,  P.  F.  Trowbridge*,  L.  D.  Haigh 


The  changes  experienced  by  beef  cattle  in  form  and  weight  and 
in  proportions  of  carcass  and  offal  when  on  different  planes  of  nu- 
trition were  presented  in  previous  bulletinsf.  The  31  representa- 
tive animals  slaughtered  at  various  intervals  from  three  groups 
were  used  (with  one  exception)  for  a study  of  the  chemical  compo- 
sition of  the  various  parts,  organs,  and  cuts  of  beef. 

GENERAL  TREATMENT 

For  a general  discussion  of  the  treatment  of  the  animals  the 
previous  bulletins  must  be  consulted.  The  ration  included  milk 
for  several  months  after  birth,  and  timothy  hay  and  grain  were 
soon  introduced.  At  weaning  time  the  ration  consisted  of  alfalfa 
hay  and  a grain  mixture  in  the  ratio  of  one  to  two.  The  grain 
consisted  of  six  parts  corn  chop,  three  parts  whole  oats,  and  one 
part  of  old  process  linseed  meal. 

The  animals  were  early  divided  into  three  groups.  Group  I 
was  fed  all  it  would  eat  of  the  ration.  Group  II  was  fed  for  maxi- 
mum growth  without  permitting  the  laying  on  of  much  fat.  Group 
III  was  fed  for  scanty  or  retarded  growth.  The  Group  II  steers 
gained  about  a pound  a day  for  the  first  two  years  while  the  Group 
III  cattle  gained  but  0.69  pounds  a day. 

The  animals  were  slaughtered  at  intervals  and  a series  of 
weights  and  measurements  were  taken.  The  wholesale  cuts  were 
divided  into  lean  flesh,  fatty  tissue,  and  bone  and  tendon.  Various 
composites  and  individual  samples  were  analyzed,  there  being  a 
rather  large  number  of  samples  for  each  animal. 

METHODS  OF  PREPARATION  OF  SAMPLES 

The  samples  of  the  soft  tissues  and  parts  were  passed  through 
a power  grinder  equipped  with  four  sets  of  plates,  each  plate  hav- 


♦Resigned  September.  191S. 

tC.  Robert  Moulton,  I*.  F.  Trowbridge,  L.  D.  Haigh,  Studies  In  Animal  Nutrition. 
1.  Changes  in  Form  and  Weight  on  Different  Planes  of  Nutrition,  Research  Bulletin 
43.  II.  Changes  in  Proportions  of  Carcass  and  Offal  on  Different  Planes  of  Nutri- 
tion, Research  Bulletin  54,  Missouri  Agricultural  Experiment  Station. 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


in g holes  of  a different  size  than  the  other.  Samples  were  ground 
through  the  coarser  plate  and  then  through  the  next  size.  The 
samples  well  mixed  and  quartered  down  if  necessary  and  then 
ground  through  a finer  plate.  The  large  samples  were  then  quartered 
again  and  ground  through  the  finest  plate.  Very  homogeneous  and  fine 
samples  were  easily  obtained  in  this  manner.  An  especially  difficult  sam- 
ple to  make  uniform  was  the  respiratory  system.  The  cartilaginous 
rings  of  the  trachea  would  partly  remain  behind  in  the  mill  while 
the  softer  lungs  were  squeezed  out  past  them.  By  means  of  a knife 
these  rings  were  finely  cut  and  mixed  with  the  lungs.  The  hide 
sample  was  cut  into  thin  strips  with  a knife,  alternate  strips  being 
rejected  in  the  larger  samples.  The  strips  were  then  cut  into  short 
lengths  and  ground  through  the  mill  already  described.  The 
grinding  of  the  sample  proceeded  very  solwly,  but  with  repeated 
grindings  the  work  advanced  more  rapidly  and  a final  uniform  and 
fine  sample  was  obtained. 

The  work  of  preparing  and  grinding  the  samples  proceeded  as 
rapidly  as  possible  until  the  samples  were  in  a position  where  there 
was  no  danger  of  decomposition  or  change.  The  samples  were 
kept  in  jars  provided  with  rubber  gaskets,  glass  tops,  and  metal 
clamps  so  that  no  loss  of  moisture  could  occur.  They  were  kept  in 
cold  storage  at  a temperature  just  above  freezing,  so  that  they  re- 
mained fresh  for  analysis. 

The  skeleton  samples  were  ground  through  a Mann  green  bone 
grinder,  mixed  well  and  sampled.  From  this  smaller  samples  were 
weighed  out  directly  and  rapidly,  in  triplicate,  in  tared  procelain 
evaporating  dishes.  The  size  of  the  samples  varied  according  to 
the  coarseness  or  fineness  of  the  bone.  For  finely  ground  samples 
25  to  40  grams  were  considered  sufficient  while  for  coarse  samples 
100  grams  or  even  more  were  sometimes  taken.  The  dishes  contain- 
ing the  weighed  samples  were  at  once  placed  in  vacuum  desiccators 
and  dried  to  a constant  weight  within  25  or  30  milligrams.  They 
were  then  extracted  with  ether  in  specially  constructed  Sohxlet 
extractors.  The  residue  was  saved,  the  triplicates  combined,  and 
the  whole  ground  in  a steel  mill  until  fine  enough  to  pass  through 
a millimeter  sieve.  The  sample  was  allowed  to  become  air  dry 
and  saved  for  a complete  analysis  later. 

Samples  of  horn  and  hoof  were  dried  and  reduced  to  a fairly 
fine  state  with  a horseshoer’s  rasp.  A drug  mill  was  then  used  to 
reduce  the  material  to  a finer  state. 


Studies  In  Animal  Nutrition — III 


5 


METHODS  OF  ANALYSIS 

The  samples  were  analyzed  for  water,  fat,  nitrogen,  ash  and 
phosphorus,  following  in  general  official  methods  of  the  Association 
of  Official  Agricultural  Chemists. 

Glycogen,  dextrose,  and  sarco-lactic  acid  and  similar  flesh 
acids  were  not  determined.  The  formation  of  the  acids  in  flesh 
progressively  increases  from  the  time  of  slaughter  up  to  a maximum 
and  then  a decrease  follows  as  decomposition  takes  place  until 
neutrality  and  finally  alkalinity  is  reached*.  The  glycogenf  con- 
tent varies  considerably  in  different  parts  of  the  animal  and  de- 
creases quite  rapidly  at  ordinary  temperatures  through  hydrolysis 
to  dextrose.  Through  determination  of  the  glycogen  content  of 
a number  of  animals  it  is  certain  that  in  beef  flesh  the  amount  of 
glycogen  will  seldom  exceed  one-half  of  one  percent. 

Water. — For  this  work  the  S.  & S.  extraction  shells  and  glass 
tubes  with  hardened  filter  paper  bottoms  were  filled  about  one-third 
full  of  ignited  sea  sand  and  then  stuffed  with  fat-free  absorbent 
cotton.  In  our  later  work  cotton  alone  was  used.  The  tubes  were 
numbered  consecutively,  extracted  with  ether,  dried  in  vacuo  and 
weighed  in  glass  stoppered  weighing  bottles.  This  was  done  pre- 
vious to  the  slaughtering.  A counterpoised  weighing  bottle  was 
found  very  convenient  as  it  obviated  complications  arising  from  a 
broken  weighing  bottle,  the  use  of  a new  bottle  and  subsequent 
corrections  of  weights.  Scheibler  vacuum  desiccators  six  inches  in 
diameter  with  stopcocks  in  the  lid  were  filled  to  the  depth  of  an 
inch  with  C.  P.  sulphuric  acid  (sp.  gr.  1.84).  A brass  gauze  or  por- 
celain plate  was  placed  on  the  shelf  of  the  desiccator  and  one-half 
inch  above  this  supported  by  corks  or  rubber  stoppers  was  a second 
gauze.  Clean  paper  was  placed  on  this.  It  was  necessary  to  have 
the  ground  glass  surfaces  and  stopcocks  fit  well.  A lubricant  of 
three  parts  of  hard  paraffin  and  five  parts  of  yellow  vaseline  was 
prepared  by  melting  together  these  ingredients  and  allowing  the 
mixture  to  cool  slowly.  In  cold  weather  a little  more  vaseline  is 
used  and  in  hot  weather  a little  more  paraffin  to  give  the  mixture 
the  proper  consistency. 

The  thoroughly  mixed  samples  were  placed  in  weighing  bottles 
provided  with  short  aluminum  scoops  and  triplicate  samples  of 
three  to  five  grams  were  weighed  out.  The  cotton  was  removed 
from  the  extraction  tube  and  placed  in  a flat-bottomed,  shallow, 


♦Trowbridge,  P.  F.  and  Orindley,  IT.  S.,  J.  Araer.  Chein.  Soc.  28,  (1906),  469. 
tTrowbridge,  P.  F.  and  Francis.  C.  K.,  J.  Ind.  and  Eng.  Chein.  2 (1910),  21  and  215. 


6 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


glazed  porcelain  dish  and  the  sand  was  poured  carefully  into  the 
dish.  The  meat  sample  was  placed  on  the  sand  and  the  whole 
was  carefully  and  thoroughly  mixed  and  then  returned  to  the  tube 
by  a steel  spatula.  The  cotton  was  used  to  wipe  every  trace  of  the 
sample  from  the  dish  and  spatula.  A large  sheet  of  glazed  paper 
prevented  loss  of  sand.  The  last  of  the  unused  cotton  was  placed 
in  the  top  of  the  tube.  Later  when  cotton  alone  was  used,  the 
mixing  of  the  sample  was  greatly  facilitated  and  the  danger  of  loss 
of  sand  was  entirely  removed.  The  sand,  or  cotton,  was  used  to  sep- 
arate the  particles  of  the  sample  and  so  allow  a more  thorough  dry- 
ing and  extraction.  Otherwise  the  samples  had  to  be  ground  and 
reextracted  a second  time.  The  triplicate  samples  were  placed  in 
separate  desiccators  in  order  to  avoid  a loss  in  case  a desiccator 
was  broken  or  acid  spilled  on  the  cones.  The  desiccators  held  15 
to  20  tubes.  The  desiccators  when  full  were  exhausted  to  a one- 
centimeter  vacuum  by  means  of  a Geryk  duplex  vacuum  pump.  The 
desiccators  were  rotated  carefully  twice  a day  to  mix  the  concen- 
trated acid  with  the  supernatant  watery  layer.  After  24  to  48  hours 
or  longer,  as  convenient,  air  was  allowed  to  bubble  slowly  through 
a sulphuric  acid  tower  into  the  desiccator  until  the  vacuum  was 
destroyed.  The  tubes  were  transferred  to  desiccators  holding  fresh 
acid  and  the  drying  was  continued  as  before.  The  tubes  were  then 
transferred  to  glass  stoppered  weighing  bottles  and  weighed  in  the 
weighing  bottle.  The  drying  was  continued  to  constant  weight 
as  given  in  detail  above. 

Fat. — The  dry  tubes  from  the  moisture  determinations  were 
extracted  for  24  hours  in  Sohxlet  extractors,  using  ether.  They 
were  partially  dried  in  an  electric  oven  at  a low  temperature  and 
then  dried  in  the  vacuum  desiccators  as  given  in  detail  above. 
They  were  weighed  as  above  and  dried  again  to  constant  weight. 
Loss  in  weight  is  fat. 

Nitrogen. — Nitrogen  was  determined  by  the  Kjeldahl-Gun- 
ning-Arnold  method.  Triplicate  samples  were  weighed  out  as  in 
the  fat  determination  and  placed  in  S.  & S.  No.  595  filter  papers 
and  introduced  into  a 500-cc.  Kjeldahl  flask.  For  hide  and  hair 
0.50  to  0.75  grams  was  used,  for  lean  meat  1.00  to  1.25  grams,  and 
for  fat  samples  2.50  to  3.50  grams.  Other  samples  in  accordance  to 
the  nitrogen  content.  Twenty-five  cubic  centimeters  of  C.  P.  con- 
centrated sulphuric  acid  was  used  for  the  meats  and  35  to  50  cc. 
for  fats.  About  0.7  grams  of  mercury  was  added  and  the  digestion 
was  made  on  a digestion  frame.  When  the  sample  had  ceased 


Studies  In  Animal  Nutrition — III 


7 


foaming  and  was  not  pasty,  7 to  10  grams  of  potassium  or  sodium 
sulphate  was  added  and  the  digestion  was  continued  for  one  or 
two  hours.  The  flasks  were  then  cooled  and  the  necks  washed 
down  with  water.  They  were  again  digested  for  an  hour  or  more. 
About  300  cc.  of  nitrogen-free  water  was  added  to  the  cool  flasks 
also  a piece  of  paraffin  the  size  of  a pea  and  a few  small  pieces  of 
granulated  zinc.  Then  85  cc.  of  the  alkali  solution  (100  cc.  for  fats) 
was  added  carefully,  the  flask  was  connected  with  a condenser,  the 
contents  were  mixed,  the  flasks  boiled  for  40  minutes  and  the  distil- 
late caught  in  a wide-mouthed  receiving  flask  containing  the  neces- 
sary amount  of  one-tenth  normal  hydrocholoric  acid  with  some 
cochineal  indicator.  The  above  alkali  solution  was  made  by  dis- 
solving 40  pounds  of  Greenbank  alkali  and  375  grams  of  potassium 
sulphide  in  30  liters  of  distilled  water.  For  fats  and  other  foam- 
ing materials  800-cc.  Kjeldahl  flasks  were  used. 

Protein. — The  protein  was  calculated  by  multiplying  the  nitro- 
gen by  the  factor  6.25. 

Ash. — Triplicate  samples  of  ten  to  fifteen  grams  were  weighed 
out  as  for  fat  and  placed  in  numbered,  tared  porcelain  crucibles. 
The  samples  were  dried  in  ovens  and  then  charred  carefully.  Later 
they  were  ashed  over  Fletcher  burners,  using  a low  heat  and  tak- 
ing plenty  of  time.  In  this  way  fusion  and  loss  of  chlorides  was 
prevented. 

Phosphorus. — The  crucibles  from  the  ash  determinations  were 
leached  with  strong  hydrochloric  acid  and  a little  nitric  acid.  The 
solutions  were  neutralized  and  ammonium  nitrate  was  added.  The 
phosphorus  was  precipitated  to  65°  C.  with  acid  ammonium  molyb- 
date. The  yellow  phospho-molybdate  was  filtered  off,  washed, 
dissolved  in  ammonia  and  hot  water  and  the  phosphorus  was  re- 
precipitated with  magnesia  mixture.  The  precipitate  was  ignited 
strongly  in  a gasoline  muffle  and  weighed  as  the  pyrophosphate. 

AIR  DRY  BONE  SAMPLES 

Moisture  and  Ash. — Two-gram  samples  were  weighed  out  in 
tared  porcelain  crucibles  and  dried  at  100  to  110°  C.  The  difference 
'in  weight  between  crucible  plus  sample  and  dry  weight  of  crucible 
plus  sample  gave  the  moisture.  The  samples  were  then  ashed  by 
igniting  over  Fletcher  burners  until  practically  free  from  carbon 
and  the  ignition  was  completed  in  a muffle  at  a dull  red  heat.  A 
clear  white  ash  was  readily  obtained  by  this  means  in  a short  time. 


8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Nitrogen. — The  nitrogen  was  determined  as  given  in  detail 
above  using  0.5  gram  samples. 

Phosphorus. — The  ash  from  the  above  determination  was  dis- 
solved by  digestion  in  hot,  dilute  nitric  acid  and  the  solution  was 
made  up  to  250  cc.  Aliquots  of  25  cc.  were  taken  and  the  phos- 
phorus determined  as  given  in  detail  above. 

COMPOSITION  OF  SAMPLES 

The  percentage  composition  of  each  sample  analyzed  is  shown 
in  the  Appendix  in  Tables  1 to  30  and  the  weights  of  the  consti- 
tuents in  Tables  31  to  60.  The  detailed  weights  for  the  separate 
parts  included  in  each  sample  can  be  found  in  Research  Bulletin 
54.  The  weight  of  the  entire  sample  is  shown  in  the  tables  listed 
above.  From  this  data  samples  can  be  composited  and  the  composition 
of  various  classes  of  tissues,  parts  of  the  animal,  or  the  entire  animal 
can  be  calculated.  The  tables  include  the  analyses  of  1061  sam- 
ples from  30  different  animals,  or  over  35  samples  per  animal. 

The  samples  listed  are  mutually  exclusive.  For  example  the 
circulatory  system  for  Steer  500  weighed  1.562  kilograms  and  had 
48.451  percent  water,  37.638  percent  fat,  and  so  on.  This  sample 
consisted  of  the  large  arteries  and  blood  vessels  in  the  thorax,  the 
pericardium,  adherent  fat,  and  the  ears  of  the  heart.  The  lean  heart 
itself  exclusive  of  the  ears  formed  a separate  sample  weighing  1.284 
kilograms  and  having  77.544  percent  of  water,  3.559  percent  of  ether 
soluble  material,  and  so  on.  Each  system  listed  is  exclusive  of 
those  parts  which  follow  as  separate  samples  which  parts  would 
ordinarily  be  considered  as  part  of  the  system. 

A few  samples  of  horns,  teeth  or  hoofs  and  dewclaws  were  lost 
or  detsroyed  before  the  analyses  were  completed.  The  composition 
of  a similar  sample  was  in  such  cases  used  to  calculate  the  compo- 
sition of  the  sample  destroyed  or  lost.  Full  explanation  of  this  is 
given  at  the  foot  of  each  table  where  such  instances  occur. 

Since  the  plan  of  the  experiment  was  slightly  modified  from 
time  to  time  the  number  and  content  of  the  samples  is  not  the  same 
for  all  the  animals.  Consequently  the  samples  can  not  all  be  com- 
pared directly.  To  facilitate  comparison  certain  composited  sys- 
tems are  presented  in  Tables  61  to  71  in  the  Appendix. 

The  Blood. — The  composition  of  the  blood  is  shown  graphi- 
cally in  figure  1.  The  water  content  is  close  to  80  percent  being 
about  82  percent  during  the  first  two  years  and  78  to  80  percent 


Studies  In  Animal  Nutrition — III 


9 


from  3 years  on.  There  is  a tendency  for  the  percentage  of  water 
to  be  in  inverse  order  to  the  plane  of  nutrition,  i.  e.,  the  higher  the 
plane  the  lower  the  percentage  of  water.  Fat  was  not  found  in  the 
blood  by  the  method  used  for  this  work. 

The  nitrogen  content  varies  between  2.5  and  3.5  percent  in- 
creasing with  age  and  increased  plane  of  nutrition,  although  there 
are  a few  exceptions  to  both  rules.  The  ash  content  is  close  to 


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Fig.  1. — Composition  of  the  blood  of  beef  animals. 


0.75  percent.  It  seems  to  be  unaffected  by  age  or  plane  of  nutri- 
tion. The  ash  content  of  the  blood  of  Steers  503,  504,  and  505  is 
unusually  low  and  probably  is  not  normal.  The  phosphorus  is 
0.025  percent  and  is  apparently  unaffected  by  the  plane  of  nutrition. 
It  seems  to  be  slightly  less  in  the  older  animals  than  in  the  younger 
animals. 

The  Nervous  System. — The  composition  of  the  central  nervous 
system — the  brain  and  spinal  cord — is  shown  in  figure  2.  The  water 
content  is  between  65  and  75  percent.  It  decreases  slightly  with 


10 


Missouri  Ac-r.  Exp.  Sta.  Research  Bulletin  55 


age  and  does  not  seem  to  be  affected  by  the  plane  of  nutrition.  The 
fat — ether  soluble  material — is  10  to  20  percent  of  this  sample.  It 
increases  with  age  and  seems  not  to  be  affected  by  the  plane  of  nu- 
trition. The  nitrogen  content  is  low  for  animal  tissue  being  about 
1.7  percent.  It  seems  to  be  independent  of  age  or  plane  of  nutri- 
tion. The  ash  varies  from  1.4  to  1.9  percent,  increasing  with  age 


Fig.  2. — Composition  of  the  central  nervous  system  of  beef  animals. 


but  being  independent  of  the  plane  of  nutrition.  The  phosphorus 
content  is  0.34  to  0.43  percent,  increasing  with  age  but  being  un- 
affected by  the  plane  of  nutrition.  Steers  503  to  505  give  too  high 
values  for  their  age.  This  sample  is  rather  typical  of  the  glands  of 
the  animal  body  being  higher  in  ash  and  phosphorus  than  any  class 
of  tissue  but  the  skeleton. 


Studies  In  Animal  Nutrition — III 


11 


Digestive  and  Excretory  System. — The  composition  of  the 
composited  digestive  and  excretory  system  is  shown  in  figure  3. 
The  external  fatty  tissue  has  largely  been  removed  from  this  sys- 
tem. The  water  is  66  to  78  percent,  the  fat  3 to  19  percent,  the 
nitrogen  2 to  2.6  percent,  the  ash  1.5  to  0.8  percent,  and  the  phos- 


Fig.  3. — Composition  of  the  digestive  and  excretory  system  of  beef 

animals. 


phorus  0.27  to  0.15  percent.  The  fat  content  increases  with  age 
and  plane  of  nutrition  while  the  water,  nitrogen,  ash  and  phos- 
phorus decrease  up  to  the  age  of  3 years.  Thereafter  the  fat  de- 
creases again  while  the  other  constituents  increase.  This  may  be 
due  to  the  fact  that  the  offal  fat  is  somewhat  more  easily  and  com- 
pletely removed  from  the  older  and  fatter  animals. 


12 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


The  Liver. — The  above  sample  is  a conglomerate  of  several 
classes  of  tissue  in  which  the  glandular  predominates.  As  an 
example  of  pure  glandular  tissue  the  liver  will  serve.  Figure  4 
shows  the  composition.  With  few  exceptions  the  composition  of 
the  liver  from  3 months  to  4 years  is  strikingly  constant.  The  wa- 
ter content  is  about  67  percent,  the  ether  soluble  matter  2 to  3 


percent,  the  nitrogen  2.8  to  3.3  percent,  the  ash  1.35  to  1.60  per- 
cent and  the  phosphorus  0.30  to  0.35  percent.  The  plane  of  nutri- 
tion does  not  seem  to  affect  the  composition,  and  age  has  but  little 
effect.  There  is  a slight  decrease  in  water  and  increase  in  fat,  nitro- 
gen and  ash  with  increasing  age.  The  ash  content  of  the  3-months- 
old  Group  I animal  and  the  phosphorus  content  of  Steer  527  are 
considered  to  be  atypical  and  are  probably  due  to  errors.  Again 


Studies  In  Animal  Nutrition — III 


13 


as  with  the  brain  and  spinal  cord  the  ash  and  phosphorus  content 
is  quite  high  while  the  nitrogen  content  is  about  that  of  muscle 
tissue. 

The  Spleen. — In  contrast  to  the  above  glands  the  spleen  (figure 
5)  has  a rather  constant  composition  and  low  fat  content.  The  fat 
runs  from  1.5  to  5 percent  and  is  slightly  greater  in  the  Group  I 
animals.  It  increases  slightly  with  age.  The  water  content  is  75 
to  78  percent.  The  nitrogen  content  is  from  2.9  to  3.3  percent  in 
the  young  animals  and  from  2.75  to  3.0  percent  in  the  old  animals. 


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ACE  IN  MONTHS. 

Fig.  5. — Composition  of  the  spleen  of  beef  animals. 


With  the  exception  of  two  of  the  Group  II  animals  the  ash  content 
is  constant  at  1.3  to  1.5  percent.  The  phosphorus  varies  from  0.260 
to  0.325  percent  in  the  young  cattle  and  from  0.300  to  0.220  percent 
in  the  old  cattle.  It  seems  to  decrease  somewhat  with  age.  On 
the  whole  the  plane  of  nutrition  has  practically  no  effect  on  the 
composition  of  the  spleen  and  there  is  but  a slight  change  with  age. 

The  Heart  and  Neck  Sweetbreads. — Not  all  of  the  other  glands 
of  the  animals  were  analyzed  as  separate  samples.  In  a number 
of  cases  the  sweetbreads,  spleen,  pancreas  and  kidneys  were  ana- 
lyzed as  separate  samples.  No  figure  is  shown  for  these  glands  but 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


a study  of  the  tables  in  the  appendix  shows  that  the  heart  and 
neck  sweetbreads  have  from  about  2 to  over  60  percent  fat  and  from 
80  to  30  percent  water.  The  nitrogen  content  runs  between  1 and 
3 percent  following  the  water  content.  The  ash  and  phosphorus 
are  1.7  and  0.34  percent  respectively  in  the  samples  low  in  fat  and 
about  one-third  those  amounts  in  the  samples  high  in  fat.  The 


fat  content  increases  in  the  older  fatter  animals  while  the  other 
constituents  decrease.  The  pancreas  is  much  like  the  heart  and 
neck  sweetbreads  in  composition.  Both  of  these  glands — the  thy- 
mus and  the  pancreas — become  so  intermingled  with  fat  in  the  older 
fatter  animals  that  a good  separation  is  impossible. 

The  Kidneys. — The  composition  of  the  kidneys  is  rather  con- 
stant. The  water  content  is  74  to  78  percent.  The  fat  content  runs 


Studies  In  Animal  Nutrition — III 


15 


from  2 to  12  percent,  but  on  account  of  the  lack  of  uniformity  in  re- 
moving the  kidney  fat  from  the  pelvis  of  the  kidney  this  variation  is 
not  considered  significant.  The  nitrogen  varies  from  2.08  to  2.7 
percent,  the  ash  from  1.00  to  1.35  percent,  and  the  phosphorus  from 
0.19  to  0.25  percent. 

The  Hair  and  Hide. — The  hair  and  hide  (figure  6)  is  a rather 
dry  tissue  having  from  50  to  70  percent  water,  1 to  13  percent  fat, 
4.8  to  6.6  percent  nitrogen,  1 to  1.5  percent  ash,  and  0.10  to  0.05 
percent  phosphorus.  The  nitrogen  content  is  higher  than  in  any 
other  tissue  excepting  hoofs,  dewclaws,  and  horn  exclusive  of  the 
bony  core.  The  fat  increases  with  age  and  plane  of  nutrition  while 
the  water  content  does  just  the  reverse.  The  nitrogen  percentage 
increases  with  age  and  is  in  inverse  order  to  the  plane  of  nutrition. 
The  ash  content  seems  to  be  rather  independent  of  age  and  nutri- 
tion. It  was  difficult  at  times  to  insure  perfectly  clean  hides  at 
slaughter  and  some  of  the  variations  in  ash  content  may  be  due 
to  dirt  on  the  animal.  The  phosphorus  content  decreases  with  age 
and  seems  to  vary  but  little  between  the  different  planes. 

The  Offal  Fat. — The  composition  of  the  offal  fat  is  shown  in 
figure  7.  A large  range  in  composition  is  shown.  This  tissue  has 
from  60  to  6 percent  of  water,  30  to  93  percent  fat,  1.7  to  0.2  percent 
nitrogen,  0.7  to  0.1  percent  ash,  and  phosphorus  0.12  to  0.01  per- 
cent. The  fat  increases  with  age  and  plane  of  nutrition,  while  all 
the  other  constituents  decrease.  The  greatest  changes  are  between 
the  ages  of  3 and  11  months. 

The  Skeleton. — The  composition  of  the  skeleton,  or  bone,  is 
shown  in  figure  8.  The  water  content  is  from  30  to  57  percent,  the 
fat  from  8 to  23  percent,  the  nitrogen  from  3 to  3.5  percent,  the  ash 
from  15  to  27  percent,  and  the  phosphorus  from  2.5  to  5 percent. 
The  fat  increases  with  age  and  is  generally  higher  in  the  well  fed 
animals  although  the  difference  is  not  great.  The  water  content  is 
just  the  reverse.  The  nitrogen  content  averages  slightly  higher 
in  the  older  animals  than  in  the  younger  animals  while  the  plane 
of  nutrition  seems  to  have  no  effect.  The  ash  and  phosphorus  con- 
tent of  the  older  animals  is  about  double  that  of  the  3-months-old 
animals.  This  ossification  is  on  the  whole  rather  gradual.  The 
plane  of  nutrition  is  here  without  effect. 

It  has  been  shown  in  earlier  work  of  this  Experiment  Station 
(Research  Bulletin  28)  that  it  is  a difficult  matter  to  alter  the  com- 
position of  the  bone  by  the  plane  of  nutrition.  The  present  study 
shows  that  aside  from  the  small  difference  in  fat  and  water  content 


16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Fig.  7. — Composition  of  the  offal  fat  of  beef  animals. 


Studies  In  Animal  Nutrition — III 


17 


Fig.  8. — Composition  of  the  skeleton  of  beef  animals. 


18 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Fig.  9. — Composition  of  the  lean  and  fat  flesh  of  beef  animals. 


Studies  In  Animal  Nutrition — III 


19 


the  bones  are  not  affected  in  composition  by  the  three  planes  of  nu- 
trition imposed. 

The  Lean  and  Fat  Flesh. — The  composition  of  the  lean  and  fat 
flesh  is  shown  in  figure  9.  This  sample  is  a composite  of  all  the 
skeletal  musculature  and  the  fatty  tissue  associated  with  it.  The 
offal  and  thoracic  fat  is  not  included.  The  figure  shows,  mainly, 
the  effect  of  increasing  fatness  on  the  composition.  The  fat  in- 
creases with  fatness  of  the  animal,  i.  e.,  with  increasing  age  and 
plane  of  nutrition,  while  all  other  constituents  decrease.  The  nitro- 
gen content  of  the  Group  II  and  Group  III  animals,  however,  is 
practically  constant  at  about  3 percent.  The  water  runs  from  77 
to  36  percent,  the  fat  from  3 to  53  percent,  the  nitrogen  from  3.2  to 
1.5  percent,  the  ash  from  1.15  to  0.50  percent,  and  the  phosphorus 
from  0.190  to  0.090  percent. 

The  Total  Animal. — The  composition  of  the  total  animal  ana- 
lyzed is  shown  in  figure  10.  The  figures  are  for  the  total  animal 
less  the  fill  and  the  loss  on  cooling  and  cutting.  This  basis  is  de- 
signated as  the  analytical  animal  in  Tables  72  and  73.  The  average 
composition  of  13  beef  calves  at  birth*  is  included  in  the  figure. 
The  water  content  decreases  from  73  percent  at  birth  to  39  percent 
in  the  old  fat  steer.  The  higher  the  plane  of  nutrition  and  the  older 
the  animal  the  lower  is  the  percent  of  water.  The  fat  increases 
from  about  4 percent  at  birth  to  about  45  percent.  The  increase 
follows  age  and  plane  of  nutrition.  The  nitrogen  shows  first  an  in- 
crease from  2.9  percent  at  birth  to  3.3  percent  at  3 months.  It  re- 
mains practically  constant  thereafter  for  Groups  II  and  III  but  de- 
creases in  Group  I to  2 percent  at  4 years.  The  ash  content  for 
Groups  II  and  III  increases  from  4.5  percent  at  birth  to  over  5 
percent  at  4 years.  The  high  value  for  the  Group  III  animal  at  40 
months  is  probably  an  error  or  an  abnormality.  For  the  Group  I 
cattle  the  ash  increases  to  about  5 percent  at  3 months,  falls  to  4 
percent  at  5J4  months  and  remains  there  in  spite  of  fattening  until 
after  3 years  when  it  drops  to  nearly  3 percent.  It  should  be  noted 
that  the  Group  III  3-months-old  calf  was  so  greatly  retarded  in 
development  by  the  low  plane  of  nutrition  that  its  ash  and  phos- 
phorus content  is  actually  lower  than  that  of  the  calves  at  birth. 
In  general  the  phosphorus  content  of  the  entire  animal  follows  the 
ash.  The  values  for  Steer  505,  Group  I,  and  Steer  503,  Group  II, 
are  so  much  higher  than  those  of  the  other  animals  of  their  groups 


Research  Bulletin  38,  Agr.  Expt.  Station,  University  of  Missouri. 


20 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


that  they  are  not  averaged  on  the  curve  but  are  shown  separately. 
The  phosphorus  content  increases  in  percentage  for  Groups  II  and 
III  but  decreases  for  Group  I. 


Studies  In  Animal  Nutrition — III 


21 


THE  COMPOSITION  ON  A PROTOPLASMIC  BASIS 

A brief  summary  of  the  composition  of  various  parts  and  sam- 
ples of  the  beef  steer  given  above  shows  that  those  parts  or  or- 
gans that  become  depots  of  deposit  for  fat,  exhibit  an  increasing 
percentage  of  fat  with  increasing  age  and  plane  of  nutrition ; while 
most,  if  not  all,  other  constituents  decrease.  Certain  organs  remain 
fairly  constant  in  composition.  It  is  the  belief  of  the  senior  author 
that  the  composition  of  animal  tissue  should  be  studied  also  on  the 
fat-free,  or  protoplasmic,  basis.  Its  usefulness  has  already  been 
demonstrated  by  Moulton*  and  Greenef.  In  the  animal  body  lipoid 
matter  can  be  divided  largely  into  two  classes:  (1)  stored  and 

inactive  lipins,  largely  glycerol  esters  of  the  higher  fatty  acids ; and, 
(2)  those  lipins  that  are  essential  to  the  protoplasmic  structure  and 
that  take  part  in  the  physiological  activities  of  the  tissue,  such  as 
lecithins  and  cholesterol.  The  former  is  largely  if  not  entirely  in- 
ert stored  matter  and  should  not  be  considered  as  part  of  the  proto- 
plasmic tissue.  Unfortunately  the  usual  method  of  extraction  by 
ether  removes  both  classes  together ; but,  since  in  the  fatty  tissue 
and  even  in  the  entire  body  of  fat  animals  the  former  very  greatly 
predominates,  the  ether  extract  can  safely  be  called  stored  fat. 

For  the  above  reasons  the  composition  will  now  be  considered 
on  the  fat-free  basis.  Tables  72  and  73  show  the  composition  of 
the  entire  animal  on  the  analytical  basis,  the  empty  weight  basis, 
and  the  fat-free  basis.  The  first  basis  has  been  defined  just  above. 
The  second  assumes  that  the  loss  on  cooling  and  cutting  is  water, 
which  it  must  largely  be.  This  weight  is  added  to  the  water  con- 
tent and  the  composition  recalculated.  The  result  is  a slightly 
larger  water  percentage  and  slightly  smaller  percentages  of  the 
other  constituents.  The  third  basis  assumes  that  all  ether  soluble 
fat  had  been  removed. 

Thirteen  calves  at  birth  and  three  embryos  reported  in  Re- 
search Bulletin  38  of  this  Station  are  included  in  the  tables.  In  or- 
der to  complete  the  picture  of  the  development  of  the  composition 
of  mammalian  tissue  a search  has  been  made  for  analyses  of  other 
mammalian  embryos.  On  account  of  the  length  of  the  gestation 
period  and  relative  size  of  the  animal  it  is  thought  that  rabbits  and 
other  mammals  cannot  serve  our  purpose.  The  composition  of 
some  21  human  embryos  reported  by  FehlingJ  in  1877  and  recalcu- 

*.T.  Biol.  Them.  XLIII,  67. 
t.T.  Biol.  Cliem.  XXXIX,  435. 
tArchlv.  f.  Gynaekologle  XI,  523. 


22 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


lated  by  the  senior  author  to  the  fat-free  basis  have  been  added  to 
the  results  obtained  from  the  bovine. 

Figure  11  shows  the  water  content  of  the  animals  on  the  fat- 
free  basis  from  the  beginning  of  gestation  to  maturity.  The  gesta- 
tion periods  for  man  and  the  ox  are  practically  the  same.  Man  is 
less  mature  at  birth,  however,  and  this  should  be  borne  in  mind  in 
considering  the  composition  of  the  full  term  human  infant.  The 
water  content  of  the  fat-free  human  embryo  at  the  beginning  of  the 
sixth  week  of  gestation,  or  at  an  intra-uterine  age  of  35  days,  is  97.5 
percent.  It  decreases  rapidly  and  uniformly  to  about  86  percent 
at  6 months.  It  is  seen  that  the  ox  embryo  at  this  age  has  practi- 
cally the  same  composition  as  the  human.  At  birth  the  ox  has  76.5 
percent  water.  The  human  infant  being  less  mature  has  81.5  per- 
cent. At  the  age  of  3 to  5 months  there  is  a marked  change  in  the 
rate  of  decrease  of  the  water  in  the  ox.  It  is  about  72  percent  at  5 
months  and  70  percent  at  4 years.  The  plane  of  nutrition  has  prac- 
tically no  effect  on  the  composition  of  the  ox,  on  the  protoplasmic 
basis. 

The  percentage  of  nitrogen  is  shown  in  figure  11.  At  about  35 
days  (intrauterine)  it  is  0.4  percent.  It  increases  rapidly  and  uni- 
formly to  about  3.0  percent  at  birth  and  at  5 months  is  3.5  percent. 
Maturity  is  reached  at  about  11  months  when  the  percentage  is 
3.6.  This  continues  to  be  the  value  excepting  for  a few  of  the  old 
thin  animals  which  exceed  it  by  about  0.2  percent. 

The  ash  content  at  35  days  is  practically  nothing.  It  increases 
rapidly  and  uniformly  to  4.3  percent  at  birth.  At  5 months  it  is  5 
percent  and  thereafter  increases  slowly  to  about  5.7  percent  at  4 
years.  There  is  more  variation  in  the  ash  content  than  in  the  wa- 
ter or  nitrogen  content.  It  is  higher  in  the  low  plane  animals  than 
in  the  high  plane  animals.  This  is  probably  due  to  a small  propor- 
tion of  bone  in  the  Group  I cattle. 

The  phosphorus  content  of  the  human  embryos  was  not  given. 
Therefore  the  figure  shows  the  results  for  the  ox  only.  The  phos- 
phorus content  on  the  protoplasmic  basis  is  about  0.3  percent  at 
185  days  intrauterine.  It  increases  rapidly  to  about  0.74  percent 
at  birth.  By  11  months  the  value  is  about  0.90  percent  and  there- 
after it  increases  very  slowly  to  1 percent  at  4 years. 

These  figures  show,  then,  that  the  evolution  of  the  tissue  of 
such  mammals  as  the  ox  is  rapid  from  conception  to  shortly  after 
birth — about  5 to  11  months  in  the  ox.  Thereafter  the  composi- 
tion on  the  fat-free,  or  protoplasmic,  basis  is  practically  constant 


Studies  In  Animal  Nutrition — III 


23 


Fig.  11. — Composition  of  the  fat-free  beef  animal. 


24 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


there  being  but  slight  changes  to  full  maturity.  Such  relations  as 
these  are  entirely  destroyed  by  the  presence  of  stored  fat  in  the 
animal  body  and  would  be  left  undiscovered  if  the  fresh,  fat-con- 
taining basis  were  used. 


THE  COMPOSITION  OF  OX  MUSCLE  ON  THE 
PROTOPLASMIC  BASIC 

The  above  results  show  the  advisability  of  studying  the  com- 
position of  such  tissues  as  striated  muscle  on  the  fat-free  basis.  Un- 
fortunately only  the  composite  embryo  was  studied  by  us.  How- 
ever, Buglia  and  Costantino*  have  recently  reported  some  analyses 
of  ox  embryo  muscle.  Three  samples  of  embryo  muscle  at  75,  120 
and  135  days  were  analyzed  for  water,  fat  and  nitrogen.  These 
results  are  included  with  all  samples  of  lean  muscle  reported  in  this 
bulletin  and  are  shown  in  Tables  74,  75,  and  76. 

Figure  12  shows  the  percentage  of  water  in  ox  muscle  from 
miduterine  life  to  maturity.  At  mid-term  the  tissue  is  87.5  percent 
water.  At  birth  it  is  80  percent  and  at  5^4  months  it  is  77  percent. 
It  remains  practically  constant  then  at  76.5  percent  with  no  appar- 
ent effect  on  the  plane  of  nutrition.  Perhaps  more  rigidly  controlled 
conditions  in  sampling  and  analyzing  might  have  resulted  in  less 
variation  than  is  shown.  The  water  content  of  the  muscle  is  5 to 
6 percent  higher  than  in  the  total  animal. 

The  nitrogen  content  is  shown  in  figure  12.  At  miduterine 
age  it  is  1.4  percent,  at  birth  2.9  percent,  and  at  11  months  3.5  per- 
cent which  is  the  value  maintained  to  the  end.  This  value  is 
slightly  less  than  that  for  the  total  animal. 

The  percentage  of  ash  exhibits  some  striking  changes.  There 
are  no  figures  preceding  birth.  At  birth  the  ash  in  the  fat-free 
muscle  is  1.05  percent.  At  3 months  of  age  it  has  risen  to  1.28  per- 
cent and  falls  to  1.11  percent  at  6 months.  From  then  on  it  de- 
creases slowly  and  gradually  to  1.06  percent  at  4 years.  The  peak 
at  3 months  may  need  verification,  but  it  is  a fact  that  only  one 
other  animal,  the  Group  II  steer  at  40  months,  exhibits  anywhere 
near  as  high  a figure. 

*Z.  Physiol.  Chemie  81  (1921),  143  and  155. 


Studies  In  Animal  Nutrition — III 


25 


The  phosphorus  percentage  in  the  fat-free  muscle  exhibits 
some  rather  similar  changes.  At  birth  the  tissue  shows  about  0.172 
percent  and  at  3 months  0.218  percent.  The  value  thereafter  falls 
fairly  rapidly  and  uniformly  to  about  0.200  percent  at  4 years.  The 
figure  confirms  in  general  the  relations  shown  by  the  ash  percent- 
ages. 


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AGE  IN  MONTHS. 

Fig.  12. — Composition  of  the  fat-free  lean  flesh  of  beef  animals. 


26 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


AMOUNT  AND  COMPOSITION  OF  GAIN 

Composition  of  Gain  From  Start  to  Slaughter. — The  compo- 
sition of  the  animal  at  slaughter  is  given  above.  In  order  to  calcu- 
late the  composition  of  the  gains  made  by  each  steer  from  the  time 
it  was  put  in  the  experiment  until  slaughter  it  is  necessary  to  know 
the  weight  and  composition  at  the  start.  The  weight  for  each  ani- 
mal at  the  beginning  of  the  experiment  is  shown  in  the  Appendix 
of  Research  Bulletin  43  of  this  series.  Since  the  analysis  is  based 
upon  empty  weight  in  Tables  72  and  73  it  is  necessary  to  estimate 
the  empty  weight  of  each  calf  at  the  start.  To  facilitate  this  the 
percentage  of  empty  weight  is  plotted  against  the  live  weight  in 


PERCENT  EMPTY  WEIGMT  IN  YOUNG 
CATTLE  AS  AFTECTEP  BY  WEIGHT 


40  50  60  70  60  90  100  110  120. 

LIVE  WEIGHT-  Kilograms. 


Fig.  13. — Percent  empty  weight  in  young  cattle. 


kilograms  in  figure  13.  Only  calves  of  100  kilograms  empty  weight 
or  less  are  shown.  The  line  shows  the  relation  between  the  per- 
centage of  empty  weight  and  the  size  of  the  animal  for  a normally 
fed  beef  calf.  The  Group  III  calves  lie  below  the  line.  The  live 
weight  used  in  this  figure  is  the  average  live  weight  for  the  last 
five  days  of  the  animal’s  life.  This  is  usually  larger  than  the  live 
weight  before  slaughter  because  at  that  time  the  cattle  had  been 
without  water  for  the  morning.  From  this  figure  it  is  possible  to 
estimate  accurately  the  probable  percentage  of  live  weight  in  each 
calf  at  the  beginning  of  the  experiment.  Table  77  gives  the  empty 
weights  at  the  start. 


Studies  In  Animal  Nutrition — III 


27 


In  figure  14  are  presented  the  relations  between  the  empty- 
weight  of  young  calves  and  the  composition  of  the  calf.  The  com- 
position at  35  kilograms  is  the  average  of  13  beef  calves  at  birth 
The  lines  show  a fairly  uniform  relation  between  empty  weight  and 
composition.  Using  the  empty  weight  of  the  calves  shown  in 
Table  77  the  composition  of  each  calf  at  the  start  can  be  accurately 
estimated. 


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EMPTY  WEIGHT  :•  Kilograms. 

Fig.  14. — Composition  of  young  cattle. 


Table  78  shows  the  weights  and  percentage  of  each  constituent 
for  each  animal  at  the  start  and  at  slaughter  and  the  composition 
of  the  gain  made.  The  animals  are  not  in  all  cases  ideal  checks 
on  each  other  consequently  the  composition  of  the  gain  does  not 
vary  uniformly.  Figure  15  shows  the  composition  of  the  gains 
made  by  each  group  from  the  start  to  slaughter. 

The  first  gains  of  the  thinnest  cattle  are  80  percent  water  the 
next  gain  is  but  62  percent  water.  The  water  increases  slightly 
to  18.5  months  and  then  decreases  slowly.  The  Group  II  cattle 


28 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Fig.  15. — Composition  of  total  gains  of  beef  cattle. 


Studies  In  Animal  Nutrition — III 


29 


show  much  the  same  sort  of  change  but  in  a less  marked  degree. 
For  the  Group  I cattle  the  gains  at  first  are  50  to  60  percent  water 
and  only  38  percent  at  4 years. 

In  contrast  to  this  the  gains  made  by  all  cattle  become  higher 
in  percentage  of  fat  as  the  age  advances.  The  Group  III  calf  at 
3 months  actually  shows  a loss  of  fat.  The  next  gain  was  10  per- 
cent fat  and  at  4 years  the  gain  contained  18  percent  fat.  The 
Group  II  cattle  show  increasing  percentages  of  fat  up  to  11  months 
when  growth  becomes  so  rapid  that  the  animal  becomes  relatively 
thinner  and  the  gains  contain  relatively  less  fat.  At  4 years  the  gain 
contains  27  percent  of  fat.  The  Group  I cattle  show  this  thinning 
down  at  8y2  months.  Thereafter  the  gains  increase  rapidly  in  fat, 
containing  at  4 years  as  much  as  46  percent  of  fat. 

The  gains  of  the  Group  I cattle  decrease  in  percentages  of 
nitrogen  excepting  during  the  period  of  rapid  growth  at  8J4  and 
11  months  when  there  is  an  increase.  At  the  start  the  gain  contains 
about  3.15  percent  nitrogen  and  at  the  end  only  1.9  percent.  The 
Group  II  cattle  show  a decrease  in  percentage  of  nitrogen  in  the 
gain  at  first  followed  by  a slight  increase.  Then  the  value  becomes 
constant  at  2.9  percent.  The  Group  III  cattle  show  much  the  same 
thing  excepting  that  the  value  continues  to  increase  up  to  40  months 
when  it  is  almost  as  great  a part  of  the  gain  as  it  was  at  3 to  5 
months. 

As  for  the  ash  gained  the  Group  III  cattle  show  a very  low  per- 
centage at  3 months  with  a very  rapid  recovery  at  5 y2  months. 
Thereafter  the  value  is  fairly  constant  at  5 percent.  The  Group  I 
and  Group  II  cattle  at  first  show  the  opposite  tendency,  the  gains 
containing  a relatively  smaller  percentage  of  ash.  The  Group  II 
cattle  then  show  a gradual  increase  to  2 years,  after  which  the  value 
is  rather  constant  at  about  5 percent.  The  Group  I cattle  continue 
to  show  a decrease  in  the  percentage  of  ash  with  some  rather  large 
individual  variations.  The  phosphorus  content  of  the  gains  made 
in  general  follows  the  ash. 

It  has  perhaps  been  noticed  that  the  lines  miss  a few  of  the 
points  by  a large  margin.  The  following  reasons  will  account  for 
this.  Steers  505  and  503,  representing  Groups  I and  II  respectively, 
were  among  the  first  animals  killed  and  analyzed.  The  other  ani- 
mals at  this  age — 11  months — differed  in  weight  or  composition 
from  these  first  two.  There  must  be  some  difference  in  age  or 
treatment  to  account  for  some  of  the  large  differences.  Steers  505 


30 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


and  503  are  not  considered  to  be  quite  typical.  Again  Steer  527, 
the  Group  I steer  at  40  months,  was  too  fat  for  its  age  and  Steers 
502  and  524  were  too  thin  for  their  ages  and  groups.  The  former 
was  the  Group  II  45-months  animal  and  the  latter  the  Group  III 
40-months  animal. 

Figure  15  (as  does  also  figure  10  which  gives  the  composition  of 
the  total  animal)  raises  the  question  of  the  regularity  of  the  change 


in  composition  of  the  cattle  and  of  the  rate  of  deposition  of  each 
constituent.  To  throw  more  light  on  these  questions  there  is  pre- 
sented in  figure  16  the  weight  of  water  found  in  each  animal 
slaughtered.  At  birth  it  is  about  25  kilograms.  In  the  Group  I 
cattle  this  increases  rapidly  and  uniformly  to  250  kilograms  at  21 
months.  The  rate  of  increase  then  declines  until  at  about  35  months 
the  steer  has  almost  as  much  water  as  at  47  months.  The  curves 


Studies  In  Animal  Nutrition — III 


31 


for  the  other  groups  are  rather  similar  excepting  that  for  Group 
II  the  break  in  the  curve  is  at  26  months  and  flattening  occurs  at 
40  months.  For  Group  III  the  rate  of  increase  has  been  still  less, 
the  break  occurs  at  about  27  months  and  the  flattening  is  at  the 
end  if  present  at  all. 

In  marked  contrast  to  these  curves  are  those  for  the  fat  shown 


in  figure  17.  After  the  third  month  the  Group  I cattle  deposited 
fat  at  a rapid  and  very  uniform  rate  averaging  7.86  kilograms  per 
month.  The  curve  is  very  slightly  convex  to  the  (abscissae)  hori- 
zontal axis.  For  the  other  groups  the  rate  of  increase  in  weight 
of  fat  is  very  much  smaller  and  the  curves  are  more  convex  to  the 
horizontal,  i.  e.,  the  rate  of  deposition  increases  more  with  age. 


32  Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 

Figure  18  shows  the  weights  of  nitrogen  (or  protein),  ash,  and 
phosphorus  for  each  animal  slaughtered.  In  general  the  curves 
resemble  the  curves  for  water  more  than  they  do  the  curves  for  fat. 
The  Group  I cattle  show  a decided  break  in  the  building  up  of  pro- 
tein at  20  months  and  a further  break  at  40  months.  The  curves 
for  the  other  groups  are  very  similar.  Both  the  ash  and  the  phos- 
phorus, on  the  other  hand,  fail  to  show  the  flattening  of  the  curve 
after  three  years. 


INCREASE.  WITH  AGE  AND  PLANE  OF  NUTRITION 


Fig.  18. — Quantity  of  nitrogen  (protein),  ash  and  phosphorus  in  beef 

cattle. 


Studies  In  Animal  Nutrition — III 


33 


COMPOSITION  OF  GAIN  BETWEEN  EACH  AGE 

In  order  to  calculate  the  composition  of  the  gains  made  be- 
tween the  succeeding  ages  it  is  necessary  to  assume  that  each  steer 
slaughtered  had  at  the  age  at  which  the  preceding  steer  was 
slaughtered  the  composition  of  that  steer  both  in  percentage  com- 
position and  in  percentage  of  empty  weight.  Table  79  gives  the 
percentage  of  empty  weight  referred  to  the  live  weight  at  the  end 
of  the  feeding  period  (a  five-day  average).  This  is  more  representa- 
tive of  conditions  in  the  pen  than  when  the  live  weight  just  preced- 
ing slaughter  is  used. 

For  the  live  weight  at  the  age  at  which  the  preceding  animal 
was  slaughtered  the  live  weight  at  the  beginning  of  that  period 
which  came  nearest  to  giving  the  correct  age  was  used.  This  fa- 
cilitates the  calculations  and  is  as  correct  as  any  of  the  assump- 
tions. The  composition  of  the  gains  made  between  each  succeeding 
age  for  each  group  is  given  in  Table  80. 

A study  of  the  table  shows  that  on  the  whole  each  animal  at 
the  time  of  slaughter  contained  more  of  each  constituent  than  it 
did  at  the  time  the  preceding  animal  was  slaughtered.  This  is  true 
with  all  but  one  animal  in  each  group  in  the  latter  months.  In  these 
cases  the  three  steers — 527  in  Group  I,  502  in  Group  II,  and  524  in 
Group  III — were  not  of  normal  condition  for  the  group,  the  first 
being  too  fat  and  the  latter  two  too  thin  for  the  age  and  group. 
These  statements  are  borne  out  by  data  presented  above  on  the 
composition  of  the  steers  and  by  the  proportion  of  lean,  fat  and 
bone  shown  by  these  animals  in  Research  Bulletin  54.  These  ani- 
mals must  be  omitted  entirely  from  a study  of  the  composition  of 
the  gains  made  between  successive  ages  or  else  the  composition 
of  normal  animals  of  the  respective  ages  and  groups  must  be  used 
in  place  of  the  composition  shown  by  those  abnormal  steers. 

From  figure  10  the  percentage  of  fat  a steer  should  have  on 
these  three  planes  of  nutrition  can  be  read  off  for  any  given  age. 
By  its  use  it  is  estimated  that  Steer  527  should  have  had  38.5  per- 
cent of  fat,  Steer  502  should  had  had  20.3  percent  of  fat,  and  Steer 
524  should  have  had  15  percent  of  fat.  From  figure  11  the  normal 
composition  of  the  fat-free  animal  is  readily  determined.  Calculat- 
ing these  values  to  the  fat  content  just  given  it  is  found  that  these 
animals  should  have  had  the  following  composition. 


34 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Fig.  19. — Composition  of  successive  gains  of  beef  cattle — water,  fat 
and  phosphorus. 


Studies  In  Animal  Nutrition — III 


35 


Estimated  Percentage  Composition. 


Steer 

Water 

Fat 

Nitrogen 

Ash 

Phosphorus 

527 

43.48 

38.5 

2.21 

3.38 

0.615 

502 

56.03 

20.3 

2.87 

4.46 

0.797 

524 

60.00 

15.0 

3.06 

4.70 

0.850 

Fig.  20. — Composition  of  successive  gains  of  beef  cattle — ash  and 
nitrogen. 


From  these  values  the  composition  of  the  gains  as  shown  in  the 
latter  part  of  the  table  are  calculated.  These  corrected  results  are 
shown  graphically  in  figures  19  and  20. 


36 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Water. — The  water  content  of  the  gains  made  by  the  Group 
I steers  decreased  as  the  animals  got  older  and  fatter.  The  8^2- 
months-old  steer  had  been  growing  rather  rapidly  and  these  gains 
were  largely  protoplasmic  tissue  as  shown  by  the  high  water  and 
nitrogen  content  of  the  gain.  From  gains  containing  60  to  70  per- 
cent of  water  the  change  becomes  rapid  to  gains  containing  30  to 
40  percent  of  water.  After  35  months  the  water  content  drops  until 
at  47  months  it  is  only  a little  over  2 percent. 

The  Group  II  steers  after  an  initial  decrease  in  percentage  of 
water  in  the  gains  show  a rise  to  11  months.  This  would  indicate 
that  their  development  was  about  3 months  behind  the  Group  I 
cattle.  At  34  months  the  steers  become  rather  fat  having  only  20 
percent  of  water  in  the  gain  made  during  the  past  8 months.  The 
next  gains  contain  more  water.  The  48-months-old  steer  of  this 
Group  was  too  fat  (and  too  well  supplied  with  bone  at  the  same 
time).  Consequently  his  last  gain  appears  to  contain  — 113  percent 
of  water.  This  is,  of  course,  impossible. 

For  the  Group  III  steers  there  is  the  initial  fall  in  percentage 
of  water  in  the  gains  followed  by  a rise  with  the  maximum  in  this 
case  deferred  to  18  months.  The  water  content  of  the  successive 
gains  then  falls  to  about  34  percent  at  45  months  and  shows  a rise 
to  about  55  percent  at  4 years. 

Fat. — In  general  the  fat  content  of  the  successive  gains  is  in- 
versely to  the  water  content.  The  first  gains  contain  about  10  per- 
cent of  fat.  This  increases,  after  an  initial  fall  with  minima  occur- 
ring where  the  water  showed  maxima,  to  90  percent  for  Group  I 
and  only  30  percent  for  Group  III.  The  Group  II  cattle  lie  between 
these,  but  on  account  of  the  wide  abnormality  of  the  4-year-old 
Group  II  steer  the  last  gain  appears  to  contain  191  percent  of  fat. 
This  is  of  course  impossible. 

Nitrogen. — The  nitrogen  content  of  the  gains  is  shown  in  fig- 
ure 20.  In  general  it  follows  the  water  and  is  inversely  to  the  per- 
centage of  fat.  The  Group  I cattle  show  gains  containing  over  3 
percent  of  nitrogen  at  first.  The  percentage  of  nitrogen  decreases 
until  at  about  4 years  the  gains  contain  no  nitrogen  ( — 0.45  and 
— 0.15  percent).  The  Group  II  and  Group  III  cattle  show  gains 
containing  over  3 percent  nitrogen  at  first.  This  percentage  falls 
to  less  than  2 percent  for  Group  III  and  about  2.5  percent  for  Group 
II.  There  is  then  an  increase  to  over  3 percent  again.  Towards 
the  end  there  are  some  rather  sudden  changes  which  can  only  be 


Studies  In  Animal  Nutrition — III 


37 


accounted  for  by  individual  differences  in  the  steers.  At  the  end 
the  values  are  not  far  from  3 percent  in  either  case. 

Ash. — The  ash  content  of  the  gains  shows  some  rather  striking 
changes.  For  the  Group  I cattle  the  percentage  drops  from  5 to 
3 or  3.6  percent.  But  the  21-months-old  steer  shows  a gain  con- 
taining over  6 percent  ash.  The  value  then  falls  to  3 percent  and 
1 percent  at  44 y2  months.  However  at  47  months  the  gains  con- 
tain nearly  7 percent  of  ash.  These  changes  are  partly  due  to  dif- 
ferent proportions  of  bone  in  the  cattle.  The  Group  II  cattle  fol- 
low Group  I in  general.  At  8^4  months  the  gain  contains  but  2.6 
percent  ash.  It  then  shows  a rise  to  nearly  6 percent  falling  rapidly 
after  26  months.  The  last  two  steers  show  abnormal  values,  the 
45-months-old  steer  showing  the  gain  of  the  last  5 months  to  con- 
tain — 3.3  percent  of  ash  and  the  4-year-old  steer  showing  for  the  last 
3 months  a gain  containing  32.37  percent  of  ash.  These  last  two 
values  are,  of  course,  impossible  and  bear  witness  to  the  abnor- 
mality of  the  4-year-old  Group  II  steer  as  well  as  to  the  different 
proportions  of  bone  in  the  last  two  animals. 

The  Group  III  animals  indicate  that  the  early  growth  of  the 
calves  has  been  so  retarded  that  the  bone  is  insufficiently  developed. 
The  gains  made  immediately  after  3 months  contain  6 percent  of 
ash  and  show  that  the  animals  are  recovering  in  this  respect.  There 
is  a big  drop  in  the  ash  content  of  the  gain  made  just  preceding  11 
months.  The  following  gain  is  higher  in  ash  and  there  then  fol- 
lows a decrease.  Towards  the  end  there  is  another  increase  in  the 
percentage  of  ash  in  the  gains. 

Phosphorus. — The  percentage  of  phosphorus  in  the  successive 
gains  is  shown  at  the  top  of  figure  19.  On  the  whole  the  values 
appear  fairly  constant.  This  is  partly  due  to  the  scale  of  the  figure. 
The  percentages  vary  much  as  do  the  ash  percentages.  The  4-year- 
old  Group  II  steer  is  again  abnormal  and  shows  the  gain  for  the 
last  3 months  to  contain  5.5  percent  of  phosphorus. 

SUMMARY 

Thirty  Hereford-Shorthorn  beef  animals  ranging  in  age  from 
3 months  to  4 years  were  used  in  this  experiment  representing  three 
different  planes  of  nutrition.  An  average  of  35  samples  per  animal 
or  a total  of  1061  samples  were  analyzed  for  water,  fat,  nitrogen, 
ash,  and  phosphorus. 

The  chief  effect  of  age  and  plane  of  nutrition  on  the  composi- 


38 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


tion  of  parts  and  total  animal  is  through  a change  in  the  fat  con- 
tent, which  increases  in  most  cases  with  age  and  plane  of  nutrition. 
The  skeleton  shows  greatly  increasing  ash  and  phosphorus  content 
with  advancing  age. 

The  total  empty  animal  shows  an  increasing  fat  content  and 
decreasing  percentage  of  other  constituents  with  age  and  plane  of 
nutrition  excepting  where  the  fattening  is  slight  and  a small  in- 
crease in  nitrogen,  ash  and  phosphorus  becomes  apparent.  When 
calculated  to  the  fat-free  basis,  however,  the  total  animal  shows 
very  striking  changes  in  composition  depending  on  age  alone.  The 
water  content  decreases  rapidly  from  conception  to  about  the  age 
of  6 months  and  then  becomes  constant.  The  other  constituents 
show  a rapid  increase  to  a maximum.  For  nitrogen  and  phosphorus 
the  maximum  is  attained  at  about  11  months.  The  ash  does  not 
attain  a maximum  and  constant  value  but  from  5 months  to  4 years 
increases  slowly. 

The  composition  of  the  composite  ox  muscle  on  the  fat-free 
or  protoplasmic  basis  shows  somewhat  similar  results.  The  mini- 
mum for  water  and  the  maximum  for  ash  occur  at  about  6 and  11 
months  respectively.  The  ash  and  phosphorus  content  show  irreg- 
ularities having  a marked  maximum  at  11  months  with  a decreasing 
percentage  thereafter. 

The  amount  and  composition  of  the  gains  from  start  to 
slaughter  and  between  each  succeeding  age  have  been  calculated. 
The  beef  steer  may  contain  4 percent  fat  at  birth  and  45  percent  at 
4 years.  For  the  full  fed  cattle  the  gains  become  richer  in  fat  and 
poorer  in  other  constituents  with  advancing  age  until  the  last  gains 
are  shown  to  consist  of  90  percent  fat.  With  the  other  groups  there 
is  some  variation.  All  groups  show  a thinning  down  during  the 
early  months  and  a fattening  after  the  period  of  rapid  growth  is 
over.  The  thin  cattle  have  a more  nearly  constant  composition 
after  the  first  few  months. 

The  irregularity  of  the  percentage  composition  of  the  gains 
raises  a question  as  to  the  uniformity  of  the  treatment.  It  is  shown 
that  the  weight  of  water  in  the  fattening  beef  steer  increases  rap- 
idly to  21  months  and  then  slowly  to  35  months.  With  the  poorer 
groups  the  flattening  of  the  curve  occurs  at  26  months  and  40 
months.  The  deposition  of  fat  was  very  uniform  from  3 months 
on  for  the  full  fed  cattle  and  slightly  increasing  with  age  for  the 
poorer  cattle. 


Studies  In  Animal  Nutrition — III 


39 


APPENDIX 


Table  1. — Steer  500.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

21,269 

1,562 

1,284 

3,747 

568 

79.041 

0.192 

3.193 

0.789 

0.022 

48.451 

37.638 

1.867 

0.541 

0.061 

77.544 

3.559 

2.723 

1.107 

0.208 

76.501 

2.702 

2.840 

1.145 

0.172 

17.038 

79.391 

0.489 

0.239 

0.024 

832 

62.766 

21.718 

1.667 

1.748 

0.371 

Digestive  and  excretory  system  (partial) 

Offal  fat 

19.275 

12.940 

538 

74.635 

13.501 

9.971 

83.556 

2.203 

0.422 

0.834 

0.184 

0.118 

0.020 

53.199 

33.943 

1.837 

1.043 

0.224 

4.634 

69.786 

2.902 

3.243 

1.579 

0.323 

Gall 

241 

91.875 

0.219 

0.200 

1.237 

0.027 

1.054 

74.374 

5.355 

2.977 

1.196 

0.219 

625 

59.139 

25.091 

2.140 

1.201 

0.256 

Kidneys 

1,019 

1,619 

77.091 

4.806 

2.420 

1.120 

0.207 

Tongue,  marketable  (excl.  bones) 

69.404 

11.874 

2.703 

0.914 

0.154 

Hair  and  hide 

35,938 

59.327 

1.319 

6.280 

1.072 

0.044 

Head  and  tail,  lean  and  fat 

3.784 

12.496 

63.713 

15.958 

3.155 

0.882 

0.134 

Shin  and  shank,  lean  and  fat 

70.862 

6.591 

3.363 

0.989 

0.164 

Flank  and  plate,  lean  and  fat 

36,410 

7,058 

58,918 

54.788 

27.651 

2.687 

0.866 

0.139 

Rump,  lean  and  fat 

55.149 

27.639 

2.527 

0.819 

0.145 

Chuck  and  neck,  lean  and  fat 

67.594 

11.869 

3.387 

0.912 

0.158 

Round,  lean 

Round,  fat 

39,898 

4,936 

29.692 

74.031 

27.767 

3.485 

61.442 

3.123 

1.590 

1.011 

0.377 

0.191 

0.051 

Loin,  lean 

70.269 

7.734 

3.113 

1.010 

0.185 

Loin,  fat 

6.830 

16.464 

76.508 

0.598 

0.245 

0.038 

Rib,  lean 

13,602 

67.137 

12.323 

3.196 

0.929 

0.170 

Rib,  fat 

1,804 

2.432 

20.368 

71.084 

1.293 

0.370 

0.060 

Kidney,  fat 

7.026 

90.275 

0.410 

0.143 

0.018 

Skeleton  of  feet 

6,838 
8 953 

39.603 

11.528 

3.612 

24.970 

4.529 

Skeleton  of  head 

47.986 

13.584 

3.487 

17.862 

3.434 

Skeleton  of  tail 

386 

39.305 

24.024 

2.650 

15.917 

2.786 

Skeleton  of  shin 

5,610 

26.487 

21.585 

3.700 

29.441 

5.211 

Skeleton  of  shank 

5,750 

31.458 

20.187 

3.454 

25.183 

4.429 

Skeleton  of  flank  and  plate 

6,350 

2,988 

14,450 

6.438 

680 

41.031 

18.008 

3.223 

18.537 

3.200 

Skeleton  of  rump 

24.341 

30.609 

3.067 

25.092 

4.430 

Skeleton  of  chuck  and  neck 

29.775 

22.517 

3.060 

25.926 

4.575 

Skeleton  of  round  (excl.  marrow) 

32.552 

27.793 

2.589 

21.074 

3.786 

Marrow  from  skeleton  of  round 

9 460 

89.251 

0.147 

0.213 

0.031 

Skeleton  of  loin 

7,772 

5.192 

25.056 

31.376 

2.935 

24.006 

4.277 

Skeleton  of  rib 

27.145 

22.308 

3.182 

27.925 

4.952 

Hoofs  and  dewclaws 

2.095 

50.581 

0.837 

7.742 

2.606 

0 117 

Teeth 

852 

21.329 

1.162 

2.079 

61.007 

11.516 

Table  2. — Steer  501.  Analysis  of  Samples. 


Description  o f sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

28,710 

77.977 

0.176 

3.290 

0.857 

0.025 

Circulatory  system 

1,836 

41.962 

45.889 

1.907 

0.476 

0.049 

Lean  Heart 

1,882 

77.616 

3.738 

2.573 

0.992 

0.198 

Respiratory  system 

3,838 

76.577 

3.348 

2.873 

1.041 

0.172 

Fat  from  tnoracic  cavity 

2,459 

18.834 

76.645 

0.612 

0.237 

0.026 

Brain  and  spinal  cord 

757 

70.404 

13.274 

1.673 

1.836 

0.392 

Digestive  and  excretory  system  (partial) 

24  235 

71.756 

12.870 

2.143 

0.809 

0.123 

Offal  fat 

38,625 

7.488 

91.061 

0.205 

0.102 

0.012 

Heart  and  neck  sweetbreads 

784 

30.756 

61.763 

0.993 

0.502 

0.107 

liver 

6,161 

69.513 

2.898 

3.233 

1.423 

0.334 

Gall 

176 

91.952 

0.050 

0.213 

1.229 

0.034 

Spleen 

1,178 

77.892 

1 953 

2.773 

1.386 

0.239 

Pancreas 

836 

59.873 

24.564 

2.203 

1.153 

0.263 

Kidneys 

1,037 

77.660 

4.867 

2.347 

1.051 

0.199 

Tongue,  marketable  (excl.  bones) 

2,153 

65.843 

16.199 

2.610 

0.870 

0.157 

Hair  and  hide 

50,090 

51.432 

13  235 

5.493 

1.522 

0.049 

Head  and  tail,  lean  and  fat 

5,224 

60.421 

20.746 

2.833 

0 767 

0 126 

Shin  and  shank,  lean  and  fat 

17,420 

58.949 

22.573 

2.707 

0.772 

0 133 

Flank  and  plate, lean  and  fat 

134,146 

26  818 

65.884 

1.050 

0.342 

0.057 

Rump,  lean  and  fat 

22.226 

28  769 

62.760 

1.140 

0.395 

0.069 

Chuck  and  neck,  lean  and  fat 

110,990 

47.698 

38.429 

1.525 

0.652 

0.118 

Round,  lean 

50,130 

69.902 

9.356 

3.090 

0.957 

0.185 

Round,  fat 

22,284 

16.846 

78.237 

0.667 

0.218 

0.026 

Loin,  lean 

45,996 

62.557 

17.934 

2.863 

0.851 

0.163 

Loin,  fat 

7 1,35  8 

9.031 

88.682 

0.388 

0.112 

0.018 

40 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  2. — Steer  501.  Analysis  of  Samples — Continued. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

20,834 

58.892 

22.409 

2.759 

0.791 

0.149 

Rib, fat 

28,322 

9.697 

87.439 

0.401 

0.134 

0.020 

Kidney,  fat 

19,544 

5.462 

93.311 

0.190 

0.067 

0.011 

Skeleton  of  feet 

7,744 

10,462 

304 

36.054 

12  326 

3.531 

26  132 

5.015 

Skeleton  of  head 

43.603 

11.776 

3.287 

23.776 

4.199 

Skeleton  of  tail 

40  880 

19.185 

3.481 

18.055 

3 155 

Skeleton  of  shin 

6,170 

7,128 

7,068 

3,682 

15,778 

6,978 

32.712 

14.188 

3.476 

28  968 

5 237 

Skeleton  of  shank 

26  943 

22.187 

3.348 

27.861 

5.124 

Skeleton  of  flank  and  plate 

40.213 

15.662 

3.320 

18.801 

3 403 

Skeleton  of  rump 

25.333 

26.213 

3.172 

25.973 

4 688 

Skeleton  of  chuck  and  neck 

30 . 106 

15.495 

3.656 

28.381 

5 205 

Skeleton  of  round  (excl.  marrow) 

26.646 

24.351 

3.086 

27.253 

4.948 

Marrow  from  skeleton  of  round 

286 

10.169 

88.390 

0.222 

0.530 

0.084 

Skeleton  of  loin 

8,614 

25.711 

22.523 

3.127 

28.411 

5.072 

Skeleton  of  rib 

6.388 

28.428 

18.371 

3.322 

27.868 

5.181 

Horns 

3,354 

36.989 

0.633 

6.469 

22.743 

4.167 

Hoofs  and  dewclaws 

2,523 

778 

47.011 

0.658 

8.453 

1.715 

0.143 

Teeth 

22.106 

0.808 

2.075 

59.784 

11.737 

Table  3. — Steer  502.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

19,728 

3,446 

77.433 

3.492 

0.720 

0.023 

Circulatory  system 

68.285 

14.513 

2.586 

0.679 

0.138 

Respiratory  system 

3,696 

76.165 

3.056 

2.829 

1.038 

0.166 

Fat  from  thoracic  cavity 

1,271 

21.504 

73.479 

0.754 

0.295 

0.041 

Brain  and  spinal  cord 

800 

69.104 

14.579 

1.717 

1.794 

0.414 

Digestive  and  excretory  system  (partial) 

20,933 

77.020 

6.035 

2.495 

1.264 

0.136 

Offal  fat 

11,377 

10.651 

86.180 

0.468 

0.185 

0.023 

Heart  andneck  sweetbreads 

502 

59.778 

25.414 

2.244 

1.153 

0.270 

Liver 

3,716 

68.941 

1.728 

3.305 

1.391 

0.334 

Gall 

241 

93.020 

0.043 

0.208 

1.027 

0.029 

Spleen 

921 

77.301 

2.420 

2.838 

1.834 

0.279 

Pancreas 

581 

53,462 

29.615 

2.172 

1.073 

0.240 

Kidneys 

838 

73.657 

6.587 

2.681 

1.115 

0.221 

Hair  and  hide 

39,556 

57.299 

3.010 

6.574 

0.976 

0.059 

Head  and  tail,  lean  and  fat 

4,250 

64.599 

14.122 

3.201 

0.826 

0.149 

Shin  and  shank,  lean  and  fat 

12,364 

68.295 

8.671 

3.486 

0.884 

0.164 

Flank  and  plate,  lean  and  fat 

35,594 

51.905 

31.222 

2.583 

0.696 

0.119 

Rump,  lean  and  fat 

8,100 

56.028 

25.426 

2.611 

0.797 

0.147 

Chuck  and  neck,  lean  and  fat 

70,744 

66.333 

12.728 

3.072 

0.921 

0.158 

Round,  lean 

44,426 

72.153 

4.051 

3.306 

0.971 

0.195 

Round,  fat 

4,620 

26.602 

62.837 

1.549 

0.322 

0.038 

Loin,  lean 

35,104 

69.877 

8.126 

3.193 

0.975 

0.199 

Loin,  fat 

9,144 

15.490 

78.182 

1.047 

0.249 

0.036 

Rib,  lean 

18,256 

66.358 

10.116 

3.068 

0.893 

0.164 

Rib,  fat 

3,338 

20.655 

69.994 

1.484 

0.281 

0.051 

Kidney,  fat 

2,916 

7.388 

89.667 

0.420 

0.241 

0.039 

Skeleton  of  feet 

6,982 

38.632 

12.205 

3.788 

23.868 

4.240 

Skeleton  of  head 

9,577 

48.757 

8.378 

3.185 

20.196 

3.460 

Skeleton  of  tail 

441 

40.425 

22.573 

3.251 

15.075 

2.733 

Skeleton  of  shin 

5,490 

27.399 

19.319 

3.532 

29.466 

5.252 

Skeleton  of  shank 

5,978 

27.197 

24.585 

3.454 

25.363 

4.505 

Skeleton  of  flank  and  plate 

5,590 

41.156 

12.997 

3.382 

21.818 

3.861 

Skeleton  of  rump 

2,362 

25.436 

29.734 

3.026 

23.684 

4.232 

Skeleton  of  chuck  and  neck 

15,092 

30.767 

18.259 

3.407 

27.553 

4.918 

Skeleton  of  round  (excl.  marrow) 

6,296 

26.477 

26.444 

2.996 

26.259 

4.681 

Marrow  from  skeleton  of  round 

408 

7.876 

90.632 

0.202 

0.413 

0.074 

Skeleton  of  loin 

7,866 

26.290 

26.514 

3.149 

25.839 

4.569 

Skeleton  of  rib 

Horns* 

5,290 
1 949 

30.433 

20.599 

3.647 

24.864 

4.401 

Hoofs  and  dewclaws 

2,010 

58.684 

0.572 

6.625 

1.186 

0.120 

Teeth 

1,038 

36.211 

1.025 

1.632 

50.004 

9.483 

•This  sample  was  lost  before  analysis. 


Studies  In  Animal  Nutrition — III 


41 


Table  4. — Steer  503.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal , 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

13,058 

82.780 

0.110 

2.708 

0.336 

0.076 

Circulatory  system 

1,782 

36.244 

54.914 

1.249 

0.329 

0.060 

Lean  heart 

1.063 

78.139 

3.736 

2.591 

0.951 

0.207 

Respiratory  system 

2,549 

78.706 

3.155 

2.654 

0.976 

0.207 

Brain  and  spinal  cord 

666 

73.106 

16.109 

1.682 

1.549 

0.394 

Digestive  and  excretory  system  (partial) .... 

8,761 

72.710 

11.013 

2.375 

1.012 

0.208 

Offal  fat 

7,385 

14.642 

81.920 

0.521 

0.182 

0.035 

Liver 

3,646 

68.680 

5.266 

2.995 

1.284 

0.334 

Kidneys 

655 

71.360 

11.795 

2.414 

1.041 

0.225 

Stomach 

5,765 

77.658 

6.925 

2.211 

1.085 

0.207 

Tongue,  marketable 

789 

69.328 

13.263 

2.535 

0.827 

0.170 

Hair  and  hide 

23,008 

67.765 

2.570 

4.793 

0.979 

0.068 

Shin,  snank,  head  and  tail,  lean  and  fat 

8.614 

69.707 

10.223 

3.196 

0.847 

0.168 

Flank  and  plate,  lean  and  fat 

16,290 

56.518 

25.666 

2.742 

0.748 

0.136 

Chuck  and  neck,  lean  and  fat 

31,934 

6S.526 

12.574 

2.954 

0.870 

0.171 

Round  and  rump,  lean 

25.022 

73.087 

4.805 

3.370 

1.024 

0.204 

Round  and  rump,  fat 

3,400 

22.450 

70.130 

1.212 

0.287 

0.048 

Loin,  lean 

17,206 

70.994 

8.451 

3.187 

0.983 

0.190 

Loin,  fat 

5.746 

15.748 

79.917 

0.777 

0.190 

0.035 

Rib,  lean 

8,932 

69.677 

10.231 

3.154 

0.927 

0.185 

Rib,  fat 

840 

23.294 

66.936 

1.481 

0.318 

0.061 

Kidney,  fat 

2,126 

8.676 

89.467 

0.340 

0.125 

0.027 

Skeleton 

41,122 

38.277 

15.059 

3.104 

23.704 

4.378 

Horns,  hoofs  and  dewclaws 

1.058 

46.286 

2.032 

7.576 

6.055 

0.661 

Teeth 

253 

23.299 

0.525 

2.252 

58.876 

11.280 

Table  5. — Steer  504.  analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

07 
/ O 

Ash 

% 

Phosphorus 

% 

Blood 

21,005 

1,637 

78.650 

3.334 

0.387 

0.405 

0.022 

0.041 

Circulatory  system 

28.710 

63.730 

1.112 

Lean  heart 

1.428 

76.650 

4.310 

2.774 

1.015 

0.203 

Respiratory  system 

3,628 

67.410 

14.900 

2.882 

0.974 

0.181 

Brain  and  spinal  cord 

724 

69.230 

15.080 

1.788 

1.358 

0.350 

Digestive  and  excretory  system  (partial) 

17,569 

68.540 

18.040 

1.910 

0.726 

0.145 

Offal  fat 

25,105 

12.760 

84.820 

0.334 

0.147 

0.023 

Liver 

4,754 

69.220 

2.436 

3.275 

1.352 

0.358 

Kidneys 

877 

69.480 

12.810 

2.452 

1.058 

0.219 

Stomachs 

12,820 

79.670 

8.040 

1.701 

0.897 

0.151 

Tongue,  marketable 

1,587 

60.750 

23.480 

2.182 

0.759 

0.141 

Hair  and  hide 

41,144 

58.290 

8.070 

5.522 

1.057 

0.043“ 

Shin,  shank,  head  and  tail,  lean  and  fat 

16,070 

60.640 

20.560 

2.951 

0.803 

0.14$ 

Flank  and  plate,  lean  and  fat 

49.650 

41.640 

45.620 

1.945 

0.572 

0.101 

Rump,  lean  and  fat 

10,846 

40.720 

46.810 

1.847 

0.574 

0.106 

Chuck  and  neck,  lean  and  fat 

59,808 

58.330 

24.030 

2.620 

0.758 

0.146 

Round,  lean 

37,238 

69.510 

9.210 

3.208 

0.983 

0.194 

Round,  fat 

9,818 

16.610 

78.030 

0.906 

0.238 

0.030 

Loin  lean 

33  676 

66.920 

12.220 

3.051 

0.946 

0.181 

Loin,  fat 

18.340 

11.620 

84.910 

0.532 

0.162 

0.025 

Rib,  lean 

18,506 

63.280 

17.520 

2.940 

0.831 

0.167 

Rib,  fat 

6,770 

14.420 

80.630 

0.833 

0.202 

0.031 

Kidney,  fat 

11,400 

4.800 

93.940 

0.215 

0.126 

0.017 

Skeleton  of  feet,  head,  tail,  shin  and  shank.. 

23,568 

36.050 

13.640 

3.237 

27.594 

5.076 

Skeleton  of  flank  and  plate 

4,572 

44 . 100 

18.180 

3.523 

15.715 

2.916 

Skeleton  of  rump 

2,428 

25.720 

26.000 

3.073 

27.381 

5.184 

Skeleton  of  chuck  and  neck 

11,176 

30.180 

16.270 

3.524 

29.162 

5.344 

Skeleton  of  round 

4,808 

21.880 

27.470 

3.120 

31.049 

5.755 

Skeleton  of  loin 

5,850 

29.840 

22.100 

3.184 

26.452 

4.812 

Skeleton  of  rib 

5.092 

32.550 

16.260 

3.417 

27.791 

5.096 

Horns,  noofsand  dewclaws 

Teeth* 

2,532 

338 

69.476 

0.514 



4.621 

2.428 

0.157 

1 

•This  sample  was  lost  before  analysis. 


42 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  6. — Steer  505.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

13,810 

82.260 

0.351 

2.726 

0.329 

0.023 

Circulatory  svstem 

1,168 

33.230 

59.064 

1.285 

0.277 

0.047 

Lean  heart 

938 

77.382 

5.110 

2.618 

0.973 

0.209 

Respiratory  system 

2,498 

76.829 

5.477 

2.713 

0.951 

0.202 

Brain  and  spinal  cord 

537 

73.810 

14.738 

1.688 

1.719 

0.411 

Digestive  and  excretory  system  (partial) 

8,258 

71.893 

12.815 

2.476 

1.053 

0.226 

Offal  fat 

12,781 

12.410 

85.384 

0.340 

0.117 

0.022 

Liver 

3,983 

68.096 

5.770 

3.205 

1.329 

0.347 

Kidneys 

718 

75.712 

7.835 

2.376 

1.061 

0.226 

Stomach 

8,818 

77.261 

10.849 

1.681 

0.868 

0.173 

Tongue,  marketable 

1,115 

64.061 

20.160 

2.352 

0.759 

0 156 

Hair  and  hide 

22,884 

62.138 

5.336 

5.297 

0.696 

0.066 

Shin,  shank,  nead  and  tail,  lean  and  fat 

9,386 

64.430 

15.398 

3.216 

0.818 

0.165 

Flank  and  plate,  lean  and  fat 

24,194 

43.720 

42.762 

2.211 

0.580 

0.116 

Chuck  and  neck,  lean  and  fat 

38,344 

62.294 

18.949 

2.886 

0.830 

0.165 

Round  and  rump,  lean 

25,784 

69.066 

9.464 

3.326 

0.976 

0.200 

Round  and  rump,  fat 

5,970 

14.140 

80.640 

0.452 

0.174 

0.032 

Loin,  lean 

19,686 

68.190 

9.983 

3.236 

0.963 

0.196 

Loin,  fat 

7,558 

9.333 

87.547 

0.535 

0.127 

0.024 

Rib,  lean 

11,300 

61.762 

19.191 

2.964 

0.841 

0.176 

Rib,  fat 

2.640 

10.913 

85.385 

0.643 

0.167 

0.033 

Kidney,  fat 

5,754 

5.263 

93.527 

0.236 

0.084 

0.016 

Skeleton 

37,745 

35.792 

17.555 

3.186 

23.852 

4.403 

Horns,  hoofs  and  dewclaws 

1,206 

46.098 

1.104 

7.724 

5.340 

0.611 

Teeth 

268 

21.938 

0.634 

2.264 

59.386 

10.697 

Table  7. — Steer  507.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

0/ 

/o 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

20,316 

78 . 174 

3.431 

0.670 

0.022 

Circulatory  system 

4,278 

47.118 

41.273 

1.695 

0.535 

0.104 

Respiratory  system 

3,768 

77.467 

3.927 

2.729 

0.954 

0.169 

Brain  and  spinal  cord 

744 

70.806 

13.738 

1.498 

1.749 

0.418 

Digestive  and  excretory  system 

27,417 

70.515 

14.079 

2.097 

0.825 

0.150 

Offal  fat 

11,037 

13.096 

83.962 

0.412 

0.140 

0.029 

Hair  and  hide 

34,473 

60.845 

6.213 

5.688 

1.085 

0.049 

Head  and  tail,  lean  and  fat 

4,038 

62.582 

18.949 

2.841 

0.895 

0.161 

Shin  and  snank.  lean  and  fat 

11.860 

69.883 

7.860 

3.343 

0.922 

0.167 

Flank  and  plate,  lean  and  fat 

36,130 

51.939 

32.362 

2.504 

0.693 

0.124 

Rump,  lean  and  fat 

7,736 

50.803 

31.644 

2.275 

0.721 

0.139 

Chuck  and  neck,  lean  and  fat 

62,530 

65.569 

15.166 

2.891 

0.862 

0.157 

Round,  lean 

39,302 

72.727 

5.772 

3.230 

0.981 

0.192 

Round,  fat 

5,378 

24.446 

68.671 

1.093 

0.276 

0.039 

Loin,  lean 

29,724 

70.696 

8.096 

3.128 

0.972 

0.185 

Loin,  fat 

10,188 

17.822 

76.684 

0.826 

0.223 

0.039 

Rib,  lean 

15,788 

67.438 

12.239 

2.999 

0.937 

0.174 

Rib,  fat 

2,432 

17.554 

76.436 

0.947 

0.253 

0.042 

Kidney, fat 

4,376 

6.784 

91.300 

0.284 

0.147 

0.025 

Skeleton  of  feet,  head  and  tail 

15,275 

42.804 

11.776 

3.456 

23.906 

3.858 

Skeleton  of  snin  and  snank 

10,350 

23.960 

19.357 

3.646 

33.717 

4.634 

Skeleton  of  flank  and  piate 

6,278 

44.217 

13.485 

3.380 

18.472 

2.851 

Skeleton  of  rump 

2,536 

25.041 

26.339 

3.229 

25.836 

3.940 

Skeleton  of  chuck  and  neck 

13,202 

31.983 

15.245 

3.555 

25.644 

4.204 

Skeleton  of  round 

5,864 

26.093 

29.961 

3.142 

23.204 

4.161 

Skeleton  of  loin 

6,506 

26.630 

26.891 

2.866 

26.102 

3.724 

Skeleton  of  rib 

5,050 

28.673 

18.021 

3.363 

28.739 

4.282 

Horns 

1,600 

41.605 

0.624 

5.762 

21.877 

3.960 

Hoofs  and  dewclaws 

1,490 

54.438 

1.143 

7.002 

2.044 

0.163 

Teetn 

712 

26.534 

1.027 

1.964 

56.717 

10.889 

Studies  In  Animal  Nutrition — III 


43 


Table  8. — Steer  509.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

18.291 

78.053 

3.429 

0.686 

0.023 

2.616 

69 . 945 

12.261 

2.590 

0.922 

0.167 

3'283 

1.248 

77.032 

2.879 

3.004 

1.011 

0.176 

19.808 

76.095 

0.720 

0.297 

0.043 

739 

66.874 

17.662 

1.679 

1 576 

0 379 

Digestive  and  excretory  svstem  (partial) 

Offal  fat 

19,016 

9.922 

630 

73.011 

11.330 

11.886 

85.888 

2.206 

0.534 

0.908 

0.175 

0.123 

0.024 

50.304 

37.140 

1.816 

1.039 

0.254 

3,875 

110 

68.408 

1.757 

3.088 

1.334 

0.326 

Gall 

91.529 

0.059 

0.250 

0.874 

0.031 

1,304 

77.255 

2.126 

2.993 

1.420 

0.242 

54.915 

28.448 

2.239 

1.178 

0.269 

774 

77.013 

3.823 

2.649 

1.115 

0.234 

37,614 

58.969 

2.477 

6.270 

1.024 

0.046 

Head  and  tail,  lean  and  fat 

3,212 

11,782 

31,790 

66.769 

13.022 

3.077 

0.821 

0 150 

Shin  and  shank,  lean  and  fat 

67.912 

9.765 

3.380 

0.911 

0.167 

Flank  and  Dlate,  lean  and  fat 

53.639 

28.926 

2.620 

0.733 

0.145 

Rump,  lean  and  fat 

7,370 

55.813 

26.599 

2.628 

0.804 

0.146 

Chuck  and  neck,  lean  and  fat 

60.176 

68.493 

9.856 

3.076 

0 895 

0.166 

Round,  lean 

40.376 

73.647 

4.183 

3.223 

1.005 

0.196 

Round,  fat 

5.106 

25.559 

64.040 

1.621 

0.343 

0.041 

Loin,  lean 

30,836 

69.632 

9.010 

3.103 

0.951 

0.180 

Loin,  fat 

7,570 

16,360 

16.259 

77.561 

1.033 

0.253 

0.041 

Rib,  lean 

67.795 

10.844 

2.927 

0 915 

0.170 

Rib,  fat 

1,978 

1,576 

6,144 

17.552 

76.022 

1.058 

0.279 

0.045 

Kidney,  fat 

5.466 

92.412 

0.314 

0.138 

0.017 

Skeleton  of  feet 

41.075 

10.778 

3.669 

23.457 

4.245 

Skeleton  of  head 

8,247 

47.540 

8.478 

3.176 

21.164 

3.772 

Skeleton  of  tail 

386 

37.947 

26.555 

2.873 

15.105 

2 . 734 

Skeleton  of  shin 

5,046 

5,498 

5,124 

2.860 

13,682 

28.351 

18.203 

3.839 

29.339 

5.168 

Skeleton  of  shank 

28.550 

22.558 

3.519 

26.144 

5.027 

Skeleton  of  dank  and  plate 

41.367 

13.032 

3.533 

22.382 

4 082 

Skeleton  of  rump 

Skeleton  of  cnuck  and  neck 

26.895 

32.387 

29.745 

17.293 

3.058 

3.704 

22.246 

25.046 

3.878 

4.606 

Skeleton  of  round  (excl.  marrow) 

5,442 

578 

27.956 

24.660 

2.967 

25.948 

4.624 

Marrow  from  skeleton  of  round 

11.658 

86.848 

0.203 

0.370 

0.058 

Skeleton  of  loin 

6.872 

27.517 

25.661 

3.130 

25.647 

4.414 

Skeleton  of  rib 

4,986 

1,590 

28.050 

16.791 

3.623 

30.569 

5.444 

Hoofs  and  dewclaws 

66.980 

0.459 

5.193 

1.457 

0.117 

Teeth 

838 

28.535 

0.535 

1.740 

56.001 

10.533 

Table  9. — Steer  512.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

24,176 

79.949 

0.055 

3.073 

0.790 

0.023 

Circulatory  system 

2.432 

40.529 

47.983 

1.710 

0.563 

0.055 

Lean  heart 

1,555 

77.321 

3.470 

2.803 

1.269 

0.215 

Respiratory  system 

3.881 

75.487 

4.670 

2.803 

1.079 

0.164 

Fat  from  thoracic  cavity 

1.171 

12.232 

85.522 

0.263 

0.144 

0.015 

Brain  and  spinal  cord 

666 

72.058 

11.117 

1.687 

1.871 

0.385 

Digestive  and  excretory  system  (partial) 

20,735 

73.684 

10.288 

2.183 

0.761 

0.119 

Offal  fat 

17,454 

11.212 

86.273 

0.369 

0.144 

0.019 

Heart  and  neck  sweetbreads 

511 

40.399 

49.421 

1.497 

0.733 

0.168 

Liver 

4.416 

68.984 

2.625 

3.263 

1.588 

0.334 

Gall 

212 

93.144 

0.033 

0.223 

1.039 

0.028 

Spleen 

1,255 

77.145 

2.366 

2.800 

1.339 

0.239 

Pancreas 

736 

57.391 

26.641 

2.087 

1.326 

0.274 

Kidneys 

1.074 

77.235 

6.831 

2.080 

1.053 

0.203 

Tongue,  marketable 

1,766 

66.379 

13.428 

2.580 

0.899 

0.161 

Hair  and  hide 

41,268 

56  193 

3.612 

6 . 547 

1.163 

0 047 

Head  and  tail,  lean  and  fat 

4.412 

61.554 

19.074 

2.893 

0.859 

0.139 

Shin  and  shank,  lean  and  fat 

12,706 

68.606 

9.380 

3.343 

0.927 

0.161 

Flank  and  plate,  lean  and  fat 

48,946 

41.914 

45.337 

1 910 

0.570 

0 094 

Rump,  lean  and  fat 

10,484 

44.598 

41.829 

2.027 

0.667 

0.119 

Chuck  and  neck,  lean  and  fat 

73,512 

63.188 

18.191 

2.826 

0.919 

0.150 

Round,  lean 

43,408 

73  272 

4.557 

3.237 

1.024 

0 192 

Round, fat 

9.940 

22.030 

70.658 

0.765 

0.311 

0 040 

Loin,  lean 

32.062 

67.607 

11.040 

3.076 

0.912 

0.170 

Loin,  fat 

15.308 

12.497 

83.354 

0.650 

0 130 

0 026 

Rib,  lean 

16,908 

65  119 

14.950 

2.967 

0.896 

0.157 

Rib,  fat 

5,398 

14.938 

80.367 

0.840 

0.212 

0.035 

44 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  9. — Steer  512.  Analysis  of  Samples — Continued. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorui 

% 

Kidney,  fat 

4,740 

4.482 

93.915 

0.183 

0.130 

0.020 

Skeleton  of  feet 

7,016 

37.452 

14.314 

3.597 

24.365 

4.405 

Skeleton  of  head 

9,665 

43.142 

12.955 

3.229 

22.420 

4.204 

Skeleton  of  tail 

416 

36.970 

24.279 

3.129 

18.107 

3.265 

Skeleton  of  shin 

6,074 

27.159 

20.808 

3.631 

29.984 

5.388 

Skeleton  of  shank 

6,156 

31.979 

21.541 

3.540 

22.768 

4.094 

Skeleton  of  flank  and  plate 

7,788 

36.792 

21.061 

3.033 

19.564 

3.586 

Skeleton  of  rump 

3,264 

23.678 

30.680 

2.986 

26.274 

4.446 

Skeleton  of  chuck  and  neck 

16.536* 

28.775 

18.986 

3.244 

30.510 

5.438 

Skeleton  of  round  (excl.  marrow) 

7,430 

28.723 

26.734 

2.822 

22.605 

4.058 

Marrow  from  skeleton  of  round 

396 

10.084 

88.297 

0.186 

0.657 

0.132 

Skeleton  of  loin 

8,748 

25.136 

24.403 

2.987 

26.642 

5.334 

Skeleton  of  rib 

6,938 

29.436 

16.324 

3.475 

28.699 

5.454 

Horns 

1,810 

35.228 

0.480 

7.033 

22.746 

3.874 

Hoofs  and  dewclaws 

1,724 

48.902 

0.588 

7.857 

2.791 

0.124 

Teeth 

710 

19.913 

0.782 

2.152 

63.721 

12.031 

Table  10. — Steer  513.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phospnorus 

% 

Blood  

25,680 

77.676 

3.341 

0.722 

0 028 

Circulatory  system 

3,485 

77.600 

4.500 

2.673 

0.909 

0.157 

Respiratory  system 

4,858 

73.905 

7.032 

2.742 

0.992 

0.155 

Fat  from  thoracic  cavity 

3,942 

13.440 

83.707 

0.366 

0.185 

0.024 

Brain  and  spinal  cord 

748 

68.192 

15.496 

1.724 

1.601 

0.410 

Digestive  and  excretory  system  (partial) 

25,642 

75.309 

7.697 

2.310 

1.041 

0.157 

Offal  fat 

53,771 

5.683 

92.874 

0.215 

0.079 

0.011 

Heart  and  neck  sweetbreads 

1,334 

40.327 

50.545 

1.405 

0.824 

0.198 

Liver 

5,920 

67.926 

3.126 

3.225 

1.397 

0.333 

Gall 

37 

91.391 

0.146 

0.312 

1.191 

0.035 

Spleen 

1,114 

75.645 

4.542 

2.932 

1.253 

0.242 

Pancreas  

873 

50.118 

34 . 657 

1.868 

1 088 

0 242 

Kidneys 

1,015 

76.071 

5.802 

2.503 

1.133 

0.220 

Hair  and  hide 

45,286 

57.566 

10.072 

5.178 

0.941 

0.054 

Head  and  tail,  lean  and  fat 

5,390 

56.379 

26.291 

2.439 

0.770 

0.128 

Shin  and  shank,  lean  and  fat 

17,798 

55.759 

27.657 

2.309 

0.697 

0.118 

Flank  and  plate,  lean  and  fat 

115,774 

29.221 

62.259 

1.242 

0.385 

0.065 

Rump,  lean  and  fat 

19,082 

31.575 

58.755 

1.378 

0.438 

0.080 

Chuck  and  neck,  lean  and  fat 

110,940 

47.879 

37.191 

2.152 

0.629 

0.116 

Round,  lean 

50,782 

65.159 

14.337 

2.901 

0.911 

0.173 

Round,  fat 

19,108 

17.763 

76.089 

0.921 

0.199 

0.024 

Loin,  lean 

44,510 

59.396 

21.597 

2.777 

0.826 

0.163 

Loin,  fat 

49,928 

8.901 

88.492 

0.366 

0.115 

0.018 

Rib,  lean 

24,744 

55.589 

27.713 

2.567 

0.757 

0.142 

Rib, fat 

23,608 

17.797 

75.455 

0.408 

0.138 

0.021 

Kidney, fat 

14,490 

3.912 

94.928 

0.156 

0.074 

0.010 

Skeleton  of  feet 

7,598 

36.238 

14.856 

3.522 

23.836 

4.243 

Skeleton  of  head 

7,865 

40.707 

9.440 

3.300 

25.028 

5.856 

Skeleton  of  tail 

297 

39.282 

19.765 

3.353 

17.252 

3.031 

Skeleton  of  shin 

6,120 

29.290 

17.835 

3.377 

27.325 

4.811 

Skeleton  of  shank 

6,058 

26.124 

19.919 

3.570 

29.591 

5.233 

Skeleton  of  flank  and  plate 

7,438 

40.922 

15.081 

3.278 

20.425 

4.139 

Skeleton  of  rump 

3,464 

24.417 

29.288 

3.009 

24.688 

4.683 

Skeleton  of  chuck  and  neck 

15,526 

32.008 

17.781 

3.508 

25.202 

5.058 

Skeleton  of  round  (excl.  marrow) 

6,564 

24.756 

30.638 

2.968 

24.757 

4.366 

Marrow  from  skeleton  of  round 

480 

8.660 

89.964 

0.161 

0.811 

0.139 

Skeleton  of  loin 

8,536 

25.186 

29.799 

3.025 

24.383 

4.699 

Skeleton  of  rib 

7,096 

26.559 

21.539 

3.363 

29.216 

5.528 

Horns 

2,144 

36.851 

0.766 

6.450 

22.622 

4.168 

Hoofs  and  dewclaws 

2,180 

41.930 

0.890 

8.956 

2.960 

0.239 

Teeth 

874 

31.897 

1.103 

1.811 

52.576 

9.959 

Studies  In  Animal  Nutrition — III 


45 


Table  11. — Steer  515.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

27,856 

5,787 

3,653 

728 

79.107 

3.217 

0.594 

0.024 

42.032 

48.530 

1.276 

0.480 

0.086 

76.354 

5.443 

2.699 

0.968 

0.187 

70.038 

16.091 

1.654 

1.634 

0.395 

36.311 

66.167 

18.711 

2.019 

1.851 

0.162 

29,877 

7.532 

90.291 

0.281 

0.124 

0.015 

49^943 

5,918 

56.003 

12.550 

4.840 

1.857 

0.059 

Head  and  tail,  lean  and  fat 

53.562 

30.538 

2.417 

0.731 

0.130 

Shin  and  shank,  lean  and  fat 

16,676 

57.361 

26.753 

2.502 

0.716 

0.123 

Flank  and  plate,  lean  and  fat 

87,138 

15,810 

31.118 

59.929 

1.360 

0.388 

0.071 

Rump,  lean  and  fat 

35.336 

54.204 

1.506 

0.476 

0.085 

Chuck  and  neck,  lean  and  fat 

88,134 

53.265 

31.088 

2.356 

0.712 

0.126 

Round,  lean 

42,942 

67.691 

11.052 

3.102 

0.924 

0.177 

Round,  fat 

19,058 

41,620 

38,324 

19,016 

17.522 

77.696 

0 784 

0.215 

0.026 

Loin,  lean  

64.691 

15.193 

2.999 

0 994 

0.181 

Loin  fat 

10.576 

86.623 

0 412 

0 115 

0.017 

Rib,  lean 

61.183 

20.387 

2.776 

0.849 

0.152 

Rib,  fat 

16.282 

9.071 

88.287 

0.394 

0.134 

0.020 

Kidney,  fat 

9,922 

4.951 

94.433 

0.178 

0.081 

0.015 

Skeleton  of  feet,  head  and  tail 

18,179 

13.900 

41.368 

11.161 

3.032 

23.490 

4.066 

Skeleton  of  shin  and  shank 

27.713 

27.063 

3.045 

24.670 

4.032 

Skeleton  of  flank  and  plate 

6,368 

4,074 

42.196 

15.964 

3.410 

17.779 

3.113 

Skeieton  of  rump 

29.793 

25 . 690 

3.096 

23.313 

4.071 

Skeleton  of  chuck  and  neck 

14,528 

6,344 

7,784 

6,464 

1.804 

30.500 

15.517 

3.333 

29.568 

4.253 

Skeleton  of  round 

23 . 643 

29.413 

3.024 

28 . 767 

4.890 

Skeleton  of  loin 

25.307 

28.219 

2.971 

26.216 

4.497 

Skeleton  of  rib 

26.086 

20.159 

3.270 

31.176 

4.374 

Horns 

40 . 640 

0.591 

5.617 

23.674 

4.223 

Hoofs  and  dewclaws 

1,893 

53.752 

0.529 

7.363 

1.823 

0.072 

Teeth 

786 

27.331 

0.859 

1.785 

57.132 

10.992 

Table  12. — Steer  523.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

15,287 

3,044 

3,371 

797 

80.520 
53  054 

2.790 

0.659 

0.598 

0.022 

Circulatory  system 

35.823 

1.684 

0.110 

Respiratory  system 

78 . 769 

4.054 

2.605 

0.980 

0 189 

Brain  and  spinal  cord 

68.595 

17.610 

1.610 

1.515 

0 359 

Digestive  and  excretory  system 

24,667 

75.194 

10.330 

2.190 

0.788 

0.158 

Hair  and  nide 

33.097 

62.801 

1.150 

5.603 

1.030 

0.051 

Head  and  tail,  lean  and  fat 

3,100 

68.166 

11.573 

3.094 

0.923 

0 . 154 

Shin  and  shank,  lean  and  fat 

8,684 

71.786 

6 038 

3.402 

0.915 

0.169 

Flank  and  plate,  lean  and  fat 

26,984 

59.510 

22 . 920 

2.693 

0.739 

0.139 

Rump,  lean  and  fat 

5.418 

54.330 

29.269 

2 532 

0.748 

0.154 

Cbuck  and  neck,  lean  and  fat 

50,320 

33,900 

25,834 

12.032 

70  884 

10  288 

2 980 

0 921 

0.171 

Round,  lean 

76.929 

2.310 

3 155 

1.042 

0.202 

Loin,  lean 

69 . 754 

10  406 

3.027 

0 921 

0.172 

Rib,  lean 

70.282 

9.297 

3.119 

0.927 

0.178 

Round,  fat 

4,556 

6,376 

1,522 

29.544 

60  224 

1.580 

0 357 

0.048 

Loin,  fat 

16.497 

77.934 

0.863 

0.281 

0.394 

0.045 

Rib,  fat 

23 . 543 

64 . 977 

1.511 

0.059 

Kidney,  fat 

3 110 

5 259 

92  726 

0 . 465 

0.179 

0.015 

Offal,  fat  

7,915 

13,120 

15.418 

44.251 

81.284 
8 120 

0 495 

0 204 

0.030 

Skeleton  of  feet,  head  and  tail 

3 563 

22.796 

4.135 

Skeleton  of  shin  and  shank 

8,862 

3.882 

29 . 464 

20  364 

3 381 

27 . 035 

5.132 

Skeleton  of  flank  and  plate  

44  868 

10.710 

3 471 

19.443 

3.467 

Skeleton  of  rump 

1,770 

9,876 

27.036 

20 . 259 

3.270 

29  591 

5.430 

8keleton  of  cbuck  and  neck 

33 . 866 

12  894 

3.681 

29.394 

5.391 

Skeleton  of  round 

4.630 

39 . 850 

26.599 

1.942 

20.522 

3 840 

Skeleton  of  loin 

5,322 

3,844 

1,167 

1,063 

766 

27.873 

23 . 686 

3.145 

27.029 

4 981 

Skeleton  of  rib 

31  218 

12  517 

3 592 

31  713 

5 769 

Horns  

46.232 

0.625 

6.616 

18.740 

3.410 

Hoofs  and  dewclaws* 

Teeth 

26.317 

0.703 

2.098 

50.395 

10.670 

•This  sample  was  lost  before  analysis. 


46 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  13. — Steer  524.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal , 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

17,019 

82.006 

2.787 

0.746 

0 022 

Circulatory  system 

2,953 

62.090 

22.418 

2.240 

0.768 

0.132 

Respiratory  system 

3,455 

77.635 

2.601 

2.793 

1.073 

0.178 

Brain  and  spinal  cord 

758 

73.218 

10.255 

1.597 

1.418 

0.338 

Digestive  and  excretory  system  (partial) 

17,924 

74.094 

9.821 

2.303 

0.885 

0.150 

Offal  fat 

5,007 

25.069 

69.444 

0.760 

0.320 

0.035 

Heart  and  neck  sweetbreads 

541 

75.123 

7.448 

2.603 

1.873 

0.463 

Liver 

3.019 

69.840 

3.316 

3.047 

1.408 

0.327 

Gall 

229 

94.859 

0.092 

0.167 

0.794 

0.031 

Spleen 

757 

78.752 

1.819 

2.750 

1.335 

0.260 

Pancreas 

435 

65.634 

15.877 

2.540 

1.230 

0.285 

Kidneys 

766 

76.830 

5.735 

2.433 

1.158 

0.208 

Hair  and  hide 

30,092 

59.259 

1.813 

6.300 

1.577 

0.058 

Head  and  tail,  lean  and  fat 

3,364 

66.261 

13.126 

3.127 

1.026 

0.179 

Shin  and  shank,  lean  and  fat 

8,674 

73.037 

5.601 

2.925 

0.974 

0.166 

Flank  and  plate,  lean  and  fat 

19,788 

63.711 

15.372 

3.200 

0.952 

0 154 

Rump,  lean  and  fat 

4,030 

63.611 

17.147 

2.957 

0.972 

0.175 

Chuck  and  neck,  lean  and  fat 

46,386 

72  555 

6 292 

3 160 

0 975 

0 174 

Round,  lean 

37,714 

76.864 

2.716 

3.257 

1.043 

0.192 

Round,  fat 

2,526 

36.260 

49.962 

2.237 

0.420 

0.053 

Loin,  lean 

24,200 

72.680 

4.544 

3.310 

1.057 

0.194 

Loin,  fat 

2,444 

18.811 

73.319 

1.237 

0.350 

0.053 

Rib,  lean  and  fat 

13,144 

70.285 

7.949 

3.310 

0.982 

0.182 

Kidney,  fat 

766 

11.440 

84.187 

0.440 

0.179 

0.031 

Skeleton  of  feet 

6,010 

40.508 

13.002 

3.592 

24.515 

4.375 

Skeleton  of  head  and  tail 

8.318 

45.930 

10.389 

3.053 

23.867 

4.263 

Skeleton  of  shin  and  shank 

10,262 

31.581 

19.317 

3.365 

25.920 

4.619 

Skeleton  of  flank  and  plate 

5,926 

43.358 

15.272 

3.211 

19.164 

3.345 

Skeleton  of  rump 

2,424 

34.304 

21.452 

2.997 

22.395 

4.057 

Skeleton  of  chuck  and  neck 

12,896 

40.299 

15.844 

3.316 

21.786 

3.986 

Skeleton  of  round 

5,878 

31.056 

28.341 

2.666 

22.700 

4.063 

Skeleton  of  loin 

6,586 

29.252 

26.901 

2.845 

24.306 

4.246 

Skeleton  of  rib 

5,310 

35.735 

18.535 

2.969 

24.951 

4.429 

Horns* 

1,227 

Hoofs  and  dewclaws 

1,494 

50.239 

0.832 

7.553 

3.194 

0.219 

Teeth 

806 

26.967 

1.106 

1.907 

57.863 

10.988 

•Tiiis  sample  was  lost  before  analysis. 


Table  14. — Steer  525.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

13,614 

80.514 

2.942 

0.660 

0.026 

Circulatory  system 

1,320 

1,658 

711 

64.087 

24.542 

1.705 

0.715 

0.121 

Respiratory  system 

78.612 

2.633 

2.823 

1.111 

0.193 

Brain  and  spinal  cord 

71.054 

13.931 

1.737 

1.504 

0.370 

Digestive  and  excretory  system 

23,001 

77.479 

9.144 

1.999 

0.883 

0.156 

Hair  and  hide 

27,813 

2,788 

7,596 

18,762 

4,154 

35,824 

62.396 

0.498 

5.724 

1.331 

0.157 

Head  and  tail,  lean  and  fat 

74.319 

13.082 

1.858 

0.986 

0.158 

Shin  and  shank,  lean  and  fat 

72.849 

5.330 

3.337 

0.965 

0.177 

Flank  amd  plate,  lean  and  fat 

60.827 

20.204 

2.873 

0.828 

0.149 

Rump,  lean  and  fat 

60 . 691 

20.925 

2.760 

0.864 

0.162 

Chuck  and  neck,  lean  and  fat 

71.151 

8.443 

3.125 

0.949 

0.178 

Round,  lean 

27,524 

18,710 

77.034 

2.403 

3.120 

1.064 

0.205 

Loin,  lean 

74.634 

3.469 

3.237 

1.048 

0.200 

Rib,  lean 

11,666 

70.514 

8.659 

3.161 

0.977 

0.181 

Round,  fat 

1,962 

3,758 

32.685 

56.826 

1.599 

0.433 

0.056 

Loin,  fat 

22.002 

69.589 

1.099 

0.294 

0.040 

Rib, fat 

664 

28.087 

61.613 

1.611 

0.605 

0.092 

Kidney,  fat 

1,258 

4,961 

10,782 

7,014 

6.721 

90.181 

0.470 

0.161 

0.020 

Offal , fat 

21.028 

70.641 

1.286 

0.293 

0.051 

Skeleton  of  feet,  head  and  tail 

43.021 

9.990 

3.736 

21.822 

4.171 

Skeleton  of  shin  and  shank 

30.333 

19.135 

3.485 

29.526 

4.190 

Skeleton  of  flank  and  plate 

3,454 

1,542 

44.613 

10.853 

3.365 

21.567 

2.989 

Skeleton  of  rump 

30.800 

23.819 

3.234 

24.612 

4.216 

Skeleton  of  chuck  and  neck 

8,450 

34 . 145 

19.714 

3.092 

22.900 

4.301 

Skeleton  of  round  

4,046 

3,928 

3,888 

28.602 

32.687 

2.653 

21.783 

2.826 

Skeleton  of  loin 

28.594 

25.309 

3.020 

24.434 

3.678 

Skeleton  of  rib 

31.735 

18.050 

3.593 

24.740 

4.352 

Horns 

1,298 

940 

48.873 

0.543 

5.641 

16.718 

3.141 

Hoofs  and  dewclaws 

51.906 

0.575 

7.892 

1.259 

0.134 

Teeth 

690 

23.125 

1.063 

1.903 

60.713 

11.737 

Studies  In  Animal  Nutrition — III 


47 


Table  15. — Steer  526.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphoru3 

% 

18,957 

79.926 

3.087 

0.777 

0.025 

2,413 

3,797 

1,585 

660 

63.316 

21.434 

2.343 

0.814 

0.145 

76.615 

3.288 

2.800 

1.025 

0.156 

21.467 

73.139 

0.753 

0.286 

0.034 

73.217 

10.255 

1.747 

1.728 

0.420 

Digestive  and  excretory  system  (partial) 

20,541 

11.551 

72.884 

12.992 

12.222 

84.144 

2.130 

0.410 

0.768 

0.175 

0.127 

0.018 

439 

60.467 

24.213 

2.253 

1.444 

0.348 

3,531 

67 . 922 

4.255 

3.247 

1.469 

0.348 

Gall  

143 

92.398 

0.138 

0.230 

1.158 

0.031 

831 

78.490 

1.496 

2.767 

1.554 

0.297 

498 

63.457 

18.209 

2.245 

1.172 

0.282 

922 

73.557 

9.071 

2.353 

1.069 

0.198 

35,732 

57.828 

5.714 

5.923 

1.440 

0.050 

3.616 

61.791 

19.640 

2.910 

0.832 

0.142 

11,644 

39,524 

70.950 

7.537 

3.301 

0 900 

0.164 

49.492 

35.249 

2.330 

0.678 

0.125 

8,594 

53.663 

29.882 

2.463 

0.738 

0.138 

61,228 

65.441 

14.500 

2 923 

0 907 

0 166 

44,614 

66.884 

11.887 

3.300 

1.031 

0.191 

5.016 

22.666 

63.981 

1.670 

0.284 

0.036 

31,440 

71.948 

5.341 

3.210 

1.018 

0.190 

11.634 

17,264 

14.508 

79.929 

0.850 

0.208 

0.030 

69.641 

10.014 

3.077 

0.920 

0.165 

3,720 

17.241 

74.768 

1.160 

0.259 

0.044 

3,224 

9.074 

87.831 

0.340 

0.137 

0.021 

6.138 

38.425 

13.853 

3.682 

23.475 

4.354 

9.165 

46.363 

12.805 

2.998 

21.004 

3.832 

11,612 

6.588 

27.802 

20.097 

3.254 

28 . 703 

5.238 

40 . 154 

16  466 

3.175 

20.333 

3 793 

2!838 

13,842 

29.344 

25  608 

3.108 

23.706 

4.277 

34.313 

17.449 

3.339 

24.601 

4.482 

6,352 

6.844 

27.057 

30.272 

2.694 

23.272 

4.318 

29.163 

28.333 

2.826 

23.314 

4.282 

Skeleton  of  rib 

5,690 

1,427 

1,875 

31.712 

20.075 

3.314 

26.287 

4.758 

Horns  

41.315 

0.744 

6.411 

19.354 

3.592 

Hoofs  and  dewclaws 

54.392 

0.625 

7.294 

2.123 

0.085 

Teeth  

782 

22.147 

1.273 

1.941 

61.470 

11.686 

Table  16. — Steer  527. 

Analysis  of  Samples. 

Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

27,382 

4.903 

78.941 

3.283 

0.762 

0.023 

Circulatory  system 

53.951 

32.798 

1.927 

0.635 

0.114 

Respiratory  system 

4,326 

3.988 

73.330 

7.186 

2.717 

1.024 

0.166 

Fat  from  thoracic  cavity 

10.340 

87.646 

0.335 

0.128 

0.015 

Brain  and  spinal  cord 

701 

69.957 

14.059 

1.583 

1.544 

0.368 

Digestive  and  excretory  system  (partial) 

Offal  fat 

23,818 

48,517 

1,037 

5,720 

1,226 

66.822 
5 . 394 

18.527 
93 . 260 

2.077 

0.183 

0.710 

0.118 

0.119 

0.013 

Heart  and  neck  sweetbreads 

36.622 

55.347 

1.187 

0.734 

0.308 

Liver 

67.872 

3 472 

3 293 

1.575 

0 184 

Spleen 

76.790 

2.169 

2.903 

1.282 

0.237 

Pancreas 

849 

41.574 

46.296 

1.460 

0.887 

0.203 

Kidneys 

1,244 

46.240 

75.345 

8.280 

2.210 

1.001 

0.193 

Hair  and  hide 

54.475 

11.859 

5.317 

1.369 

0.056 

Head  and  tail.  lean  and  fat 

5,018 

53.921 

30 . 635 

2.410 

0.691 

0.123 

Shin  and  shank,  lean  and  fat 

17,358 

56.503 

26.398 

2.570 

0.754 

0.130 

Flank  and  plate,  lean  and  fat 

118.978 

27.200 

65.491 

1.097 

0.312 

0.059 

Rump,  lean  and  fat 

24,020 

29  026 

62.962 

1.233 

0.315 

0.059 

Chuck  and  neck,  lean  and  fat 

112.440 

46.782 

39.316 

2.030 

0.617 

0.109 

Round,  lean 

51.396 

66.109 

13.931 

2.993 

0.864 

0.175 

Round,  fat 

21,466 

50.140 

16.129 

79.323 

0.753 

0.204 

0.022 

Loin,  lean 

61  192 

20.496 

2.797 

0.849 

0.161 

Loin,  fat 

52,724 

9.519 

88.271 

0.357 

0.127 

0.017 

Rib,  lean 

25,860 

54 . 984 

28.098 

2.547 

0.737 

0.141 

Rib,  fat 

24.278 

9.463 

88 . 208 

0.407 

0.133 

0.016 

Kidney,  fat 

18.964 

5.423 

93.227 

0.187 

0.102 

0.014 

Skeleton  of  feet  

7,442 
8 822 

37.280 

16.115 

3.372 

23 . 664 

3.849 

Skeleton  of  head  and  tail 

42  136 

24.822 

3.165 

21.749 

3.076 

Skeleton  of  shin  and  shank 

13,136 

6,082 

27 . 256 

21 . 276 

3.149 

28.369 

4 550 

Skeleton  of  flank  and  plate  

39.334 

18.437 

2.996 

19.531 

3.404 

Skeleton  of  rump  

3.260 

24  846 

30 . 690 

2.967 

23.925 

3.747 

Skeleton  of  chuck  and  neck 

14,870 

28.058 

24.570 

3.082 

25.037 

3.944 

Skeleton  of  round 

6,446 

20.998 

33 . 954 

2.642 

27.032 

4.964 

Skeleton  of  loin 

7,140 

25.920 

25.782 

3.151 

20.949 

5.156 

Skeleton  of  rib 

6,546 

26.920 

23.074 

3.022 

28 . 622 

5.503 

Homs 

1,266 

35 . 662 

0.775 

6.950 

19.811 

3 . 658 

Hoofs  and  dewclaws 

2.174 

44.082 

0.959 

8.916 

2.059 

0.145 

Teeth 

872 

20.697 

1.438 

1.954 

63.378 

11.767 

48 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  17. — Steer  531.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

ABh 

% 

Phosphorus 

% 

9.457 

81.882 

2.841 

0.703 

0.040 

0.126 

0.204 

0.363 

0.121 

0.046 

0.427 

0.321 

0.052 

0.291 

0.288 

0.213 

0.075 

0.155 

0.188 

0.176 

0.177 

1,971 

56.721 

29.318 

1.968 

0.676 

1,915 

573 

78.338 

2.818 

2.680 

1.054 

72.987 

13  498 

1.765 

1 502 

Digestive  and  excretory  system  (partial) 

Offal  fat 

11,807 

2,899 

77.129 

25.931 

9.230 

70.680 

1.977 

0.621 

0.669 

0.290 

472 

67.988 

14.791 

2.497 

1.720 

2,205 

86 

71.785 

1.945 

2.921 

1 385 

Gall 

91.678 

0.214 

0 978 

481 

75.079 

4.195 

2.958 

1 440 

Pancreas 

297 

67.078 

14.763 

2.508 

1.229 

Kidneys  

506 

75.079 

8.283 

2.265 

1.042 

Hair  and  nide 

16.693 

63.665 

0.811 

5.733 

1.136 

Head  and  tail,  lean  and  fat 

1,722 

5.480 

10.854 

67.218 

14.073 

2.923 

0 872 

Shin  and  shank,  lean  and  fat 

71.792 

4.736 

3.596 

1 071 

Flank  and  plate,  lean  and  fat 

60.374 

17.191 

3.055 

1.005 

Rump,  lean  and  fat 

2,506 

61.990 

18.113 

2.980 

0.976 

Chuck  and  neck,  lean  and  fat 

26,902 

21,496 

1,490 

14078 

69.914 

7.805 

3.274 

1.034 

0 196 

Round,  lean 

75.102 

1.831 

3.272 

1.102 

0.208 

0.071 

Round,  fat 

28.477 

60.684 

1.524 

0.478 

Loin,  lean 

73.011 

4.278 

3.312 

1.100 

0.204 

Loin,  fat 

2.264 

24.756 

64.861 

1.386 

0.439 

0.075 

Rib,  lean  and  fat 

6 612 

69.536 

7.559 

3.338 

1.052 

0.190 

Kidney  fat 

726 

6.066 

90.265 

0.596 

0.240 

0.031 

Skeleton  of  feet 

3,762 

4.842 

39.773 

14.354 

3.233 

24.642 

4.432 

Skeleton  of  head  and  tail 

51.145 

8.071 

3.041 

20.849 

3.629 

Skeleton  of  shin  and  shank 

5.998 

31.730 

18.346 

2.937 

27.416 

5.113 

Skeleton  of  flank  and  plate 

2,546 

46.771 

10.571 

3.220 

20.395 

3.464 

Skeleton  of  rump 

1.060 

31.418 

21.238 

3.274 

25.533 

4.573 

Skeleton  of  chuck  and  neck 

6,098 

37.493 

16.015 

3.536 

23 . 526 

4.187 

Skeleton  of  round 

3.640 

35.030 

25.913 

2.679 

21.363 

3 806 

Skeleton  of  loin 

2.834 

31.090 

21.915 

3.298 

25.494 

4 471 

Skeleton  of  rib 

2.280 

32.384 

16.742 

3.738 

25.229 

4 587 

Hoofs  and  dewclaws 

760 

52 . 103 

0.827 

7.587 

1.963 

0.125 

Teeth 

426 

27.423 

0.882 

1.990 

55.799 

10.625 

Table  18. — Steer  532.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

18,752 

4,483 

80.473 

2.997 

0.652 

0.033 

Circulatory  system 

47.915 

40.256 

1.605 

0.566 

0.105 

Respiratory  system 

3,870 

77.557 

3.413 

2.636 

1.052 

0.193 

Brain  and  spinal  cord 

643 

72.457 

14.458 

1.658 

1.622 

0.392 

Digestive  and  excretory  system  (partial) 

Offal  fat 

21,741 

23,697 

441 

72.701 

9.552 

13.979 

88.805 

1.898 

0.243 

0.708 

0.116 

0.134 
0 016 

Heart  and  neck  sweetbreads 

49.463 

38.461 

1.629 

0.986 

0.238 

Liver 

5,694 

185 

71.419 

2.375 

2.843 

1.375 

0.307 

Gali 

92  254 

0.220 

1.005 

0.043 

Spleen 

884 

74.698 

5.033 

2.888 

1.256 

0.262 

Pancreas 

630 

56. 164 

27.676 

2.032 

1.180 

0.293 

Kidneys 

868 

71.247 

10.983 

2.521 

1.019 

0.213 

Hair  and  hide 

33,988 

4,260 

59.564 

6.839 

5.225 

1.032 

0.071 

Head  and  tail,  lean  and  fat 

60.370 

21.950 

2.687 

0.807 

0.146 

Shin  and  shank,  lean  and  fat 

12,008 

67.536 

10.998 

3.180 

0.933 

0.167 

Flank  and  plate,  lean  and  fat 

44,636 

43.753 

41.749 

2.237 

0.655 

0.116 

Rump,  lean  and  fat 

8,058 

48.528 

35.883 

2.345 

0.716 

0.133 

Chuck  and  neck,  lean  and  fat 

66.204 

61.792 

18.520 

2.910 

0.899 

0.156 

Round,  lean 

38.064 

71.904 

5.223 

3.304 

1.065 

0.200 

Round,  fat 

6.064 

21.714 

70.321 

1.113 

0.295 

0.042 

Loin,  lean 

36,136 

68.406 

9.593 

3.252 

1.004 

0.184 

Loin,  fat 

14,954 

12.520 

83.405 

0.674 

0.191 

0.033 

Rib,  lean 

17,356 

67.280 

11.483 

3.156 

0.979 

0.176 

Rib,  fat 

6.194 

15.622 

78.878 

0.852 

0.268 

0.040 

Kidney  fat 

11.734 

<5.271 

95.229 

0.150 

0.085 

0.022 

Skeieton  of  feet 

6.490 

39.080 

14.349 

3.607 

23.845 

4.221 

Skeleton  of  head  and  tail 

7.120 

46.409 

12.787 

2.985 

21.246 

3.794 

Skeleton  of  shin  and  shank 

10.756 

28.890 

21.617 

4.316 

23.357 

4.397 

Skeleton  of  flank  and  plate 

5.478 

44.408 

18.946 

3.027 

14.958 

2.738 

Skeleton  of  rump 

2.282 

28.899 

27.742 

3.205 

22.689 

4.084 

Skeleton  of  chuck  and  neck 

13.014 

30.283 

23.624 

3.367 

24.027 

4.351 

Skeleton  of  round 

5.624 

26.867 

34.032 

2.514 

20.794 

3.713 

Skeleton  of  loin 

6.246 

28.097 

29.257 

3.093 

22.053 

4.074 

Skeleton  of  rib 

5 222 

34.969 

22.509 

2.951 

22.280 

4.009 

Horns 

228 

54.753 

0.511 

6.327 

7.572 

1.439 

Hoofs  and  dewclaws 

1.406 

494 

51.936 

0.646 

7.653 

1.948 

0.153 

Teeth 

31.017 

0.882 

2.145 

51.709 

10.154 

Studies  In  Animal  Nutrition — III 


49 


Table  19. — Steer  538.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

7,219 

1.670 

82.400 

0.097 

2.735 

0.786 

0.028 

55.482 

28.703 

1.934 

0.632 

0.114 

1,825 

80.802 

2.057 

2.503 

1.018 

0.199 

4S7 

74.331 

11.750 

1.610 

1.500 

0.348 

Digestive  and  excretory  system  (partial) 

10,290 

3,452 

481 

76.022 

21.182 

9.813 

76.453 

2.111 

0.544 

0.867 

0.272 

0.151 

0.055 

69.816 

14.598 

2.329 

1 . 655 

0.405 

1,978 

199 

70.479 

1.914 

2.958 

1.402 

0.315 

Gall 

90.916 

0.078 

0.224 

1.038 

0.070 

331 

78.439 

1.530 

2.877 

1.409 

0.272 

208 

69.655 

11.646 

2.566 

1.465 

0.353 

487 

73.220 

11.270 

2.305 

1.075 

0 209 

15.342 

1,496 

64.339 

1.342 

5.564 

1.063 

0 063 

Head  and  tail,  lean  and  fat 

66.304 

15.440 

2.701 

0.893 

0.148 

Shin  and  snank,  lean  and  fat 

4,190 

11.036 

71  811 

5.773 

3.294 

1.002 

0.179 

Flank  and  plate,  lean  and  fat 

57  700 

23.466 

2.774 

0.852 

0.160 

Rump,  lean  and  fat 

2,096 

59  745 

21.117 

2.669 

0.892 

0 154 

Chuck  and  neck,  lean  and  fat 

22.284 

68.487 

11  963 

2.971 

0.931 

0 174 

Round,  lean 

16.324 

75.971 

2.699 

3.159 

1.093 

0.202 

Round,  fat 

1,702 

12.732 

30.528 

58.733 

1.707 

0.449 

0.068 

Loin,  lean 

73.141 

5.617 

3.117 

1.055 

0.203 

Loin,  fat  

2,360 

5,196 

19.936 

72.602 

0.977 

0 293 

0.054 

Rib.  lean 

70.881 

8.150 

3.075 

1.017 

0.191 

Rib.  fat 

426 

30 . 179 

55.423 

1.454 

0.725 

0.114 

Kidney,  fat 

622 

6.747 

90 . 964 

0.340 

0.176 

0.034 

Skeleton  of  feet 

3.168 

42 . 658 

15.526 

3.293 

18.615 

3.342 

Skeleton  of  head 

4,001 

49.779 

7.792 

2.917 

21.579 

4.089 

Skeleton  of  tail 

135 

52  455 

14.133 

3.189 

11  275 

1 944 

Skeleton  of  shin 

2.208 

28  636 

23  827 

2.827 

25  111 

3 924 

Skeleton  of  shank 

2.608 

30.367 

22.437 

3.389 

23.653 

3.670 

Skeleton  of  flank  and  plate 

2,144 

49.188 

14.542 

3.198 

15.045 

2.447 

Skeleton  of  rump 

732 

34.282 

23.745 

3.050 

22.231 

3.797 

Skeleton  of  chuck  and  neck 

5.412 

38.139 

16.952 

3.337 

23.969 

3.555 

Skeleton  of  round 

2.556 

29.961 

32.225 

2.423 

20 . 600 

3.177 

Skeleton  of  loin 

2.696 

34 . 968 

23.196 

2.940 

21.259 

4.163 

Skeleton  of  rib 

2,154 

36.366 

16.613 

3.179 

22.619 

4.031 

Horns 

250 

54 . 908 

0.529 

5.783 

9.976 

1.895 

Hoofs  and  dewclaws 

635 

66  631* 

0.465 

5.359 

1.015 

0.067 

Teeth 

240 

28.678f 

1.111 

1.961 

54.175 

10.200 

*Hoofs  and  dewclaws  of  steer  540  and  steer  538  were  analyzed  together, 
t Teeth  of  steer  540  and  steer  538  were  analyzed  together. 


Table  20. — Steer  540.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

6,967 

1,436 

1.501 

82.280 

1.030 

2.806 

0.724 

0.026 

Circulatory  system 

58.131 

27.726 

2.039 

0.676 

0.127 

Respiratory  system 

79.609 

2.314 

2.592 

1.032 

0.198 

Brain  and  spinal  cord 

512 

73.656 

11.940 

1.617 

1.579 

0.375 

Digestive  and  excretory  system  (partial) 

Offal  fat 

9,280 

2,307 

408 

75.239 

25.244 

10.756 

71.058 

1.995 

0.548 

1.703 

0.282 

0.129 

0.047 

Heart  and  neck  sweetbreads 

68.793 

15.449 

2.258 

1.510 

0.373 

Liver 

1,593 

70.592 

1.760 

2.840 

1.411 

0 . 305 

Gall 

58 

93.842 

0.076 

0.168 

1.224 

0.040 

Spleen 

331 

77.793 

1.372 

3.097 

1.408 

0.271 

Pancreas 

180 

72.201 

9 . 605 

2.537 

1.457 

0.343 

Kidneys 

363 

72.153 

10.499 

2.368 

1.084 

0.223 

Hair  and  hide 

12,994 

64 . 642 

2.353 

5.127 

1.256 

0.067 

Head  and  tail,  lean  and  fat.  . . 

1,274 

07  992 

13.404 

2.736 

0.883 

0.167 

8hin  and  shank,  lean  and  fat 

3,762 

74 . 264 

4.038 

3.397 

1 .039 

0 185 

Flank  and  plate,  lean  and  fat 

8,824 

60.923 

18.461 

3.021 

0.883 

0.158 

Rump,  lean  and  fat 

1,964 

01  016 

19  592 

2.755 

0.920 

0.166 

Chuck  and  neck,  lean  and  fat 

17,978 

71  ins 

8.347 

2.961 

0.986 

0.178 

Round,  lean 

13.456 

75 . 698 

2.255 

3.208 

1.082 

0.210 

Round,  fat 

810 

27.830 

62.313 

1.356 

0.388 

0.060 

Loin,  lean 

10,700 

73  645 

4.187 

3.179 

1.075 

0.201 

Loin,  fat 

2.308 

10  598 

73.692 

0 . 958 

0.288 

0.058 

Rib,  lean  and  fat 

5,046 

69  429 

9 677 

3.096 

1.013 

0.181 

Kidney,  fat 

682 

13.401 

79.809 

1.156 

0.185 

0.035 

50 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  20. — Steer  540.  Analysis  of  Samples — Continued. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Skeleton  of  feet 

2,784 

3,682 

44.008 

12.128 

3.288 

20.841 

3.573 

Skeleton  of  head 

51.688 

7.055 

2.754 

21.417 

4.056 

Skeleton  of  tail 

138 

55.385 

12.477 

2.254 

10.059 

1.771 

Skeleton  of  shin 

1,952 

31.687 

22.158 

3.498 

24.179 

4.561 

Skeleton  of  shank 

2,304 

1,862 

33.134 

24.266 

2.969 

21.100 

3.760 

Skeleton  of  flank  and  plate 

53 . 192 

12.085 

3.378 

12.087 

2.902 

Skeleton  of  rump 

622 

35.209 

20.449 

3.031 

22.451 

4.221 

Skeleton  of  chuck  and  neck 

4.896 

41.285 

14.741 

3.264 

19.664 

3.532 

Skeleton  of  round 

2,350 

2.550 

35.287 

27.370 

2.523 

16.708 

2.801 

Skeleton  of  loin 

34.808 

27.381 

2.786 

19.208 

3.575 

Skeleton  of  rib 

1.824 

34.170 

18.773 

3.297 

22.860 

4.637 

Horns  

304 

54  266 

0.593 

5.453 

12.472 

2.411 

Hoofs  and  dewclaws 

635 

66.631* 

0.465 

5.359 

1.015 

0.067 

Teeth 

278 

28 . 678f 

1.111 

1.961 

54.175 

10.200 

*Hoofs  and  dewclaws  of  steer  538  and  steer  540  were  analyzed  together. 
fTeeth  of  steer  538  and  steer  540  were  analyzed  together. 


Table  21. — Steer  541.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal , 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

C7 

/o 

Blood 

12,470 

2.646 

2,387 

568 

82.112 

0.089 

2.773 

0.679 

0.027 

Circulatory  system 

48.463 

39.878 

1.648 

0.572 

0.105 

Respiratory  system 

79.217 

1.976 

2.738 

1.040 

0.213 

Brain  and  spinal  cord 

72.458 

13.605 

1.658 

1.525 

0.371 

Digestive  and  excretory  system  (partial) .... 
Offal  fat 

16.313 

11,009 

740 

71.285 

11.714 

14.700 

86.398 

2.024 

0.293 

0.746 

0.125 

0.134 

0.026 

Heart  and  neck  sweetbreads 

65.439 

19.148 

2.280 

1.457 

0.367 

Liver 

3,832 

202 

69.540 

2.502 

3.158 

1.441 

0.349 

Gall 

89.007 

0.174 

0.286 

1.207 

0.060 

Spleen 

596 

77.687 

1.811 

2.876 

1.326 

0.279 

Pancreas 

390 

59.915 

23.448 

2.036 

1.192 

0.285 

Kidneys 

645 

73.472 

9.902 

2.223 

1.076 

0.214 

Hair  and  hide 

26,574 

2,062 

6,830 

24,910 

4,454 

40,480 

27,000 

3,854 

23,416 

60.425 

3.650 

5.692 

1.100 

0.055 

Head  and  tail,  lean  and  fat 

66.470 

14.520 

2.939 

1.016 

0.164 

Shin  and  shank,  lean  and  fat 

68.724 

9.020 

2.923 

0.956 

0.178 

Flank  and  plate,  lean  and  fat 

49.802 

34.241 

2.427 

0.717 

0.127 

Rump,  lean  and  fat 

50.532 

32.775 

2.370 

0.713 

0.137 

Chuck  and  neck,  lean  and  fat 

65.263 

14.772 

2.942 

0.913 

0.165 

Round,  lean 

73.915 

3.366 

3.340 

1.089 

0.208 

Round,  fat 

21.202 

73.399 

0.987 

0.282 

0.046 

Loin,  lean 

71.952 

5.559 

3.233 

1.050 

0.198 

Loin,  fat 

8.088 

14.861 

80.634 

0.780 

0.204 

0.039 

Rib,  lean 

11,588 

2,434 

68.657 

10.433 

3.126 

0.796 

0.187 

Rib, fat 

17.784 

76.039 

1.080 

0.312 

0.059 

Kidney, fat 

6,056 

4,506 

5,400 

4.494 

94.350 

0.202 

0.086 

0.020 

Skeleton  of  feet 

41.552 

12.840 

3.435 

21.608 

3.743 

Skeleton  of  head 

50.560 

7.061 

2.995 

20.107 

3.657 

Skeleton  of  tail 

206 

47.886 

19.594 

3.175 

10.884 

1.894 

Skeleton  of  shin 

2,946 

3,744 

2,864 

1,056 

7,296 

28.889 

23.422 

3.102 

27.138 

3.798 

Skeleton  of  shank 

32.657 

20.792 

2.933 

27.763 

3.778 

Skeleton  of  flank  and  plate 

51.920 

15.329 

2.277 

11.840 

1.799 

Skeleton  of  rump 

30.290 

24.456 

3.061 

25.444 

4.853 

Skeleton  of  chuck  and  neck 

34.849 

18.389 

3.368 

25.110 

3.622 

Skeleton  of  round 

3,250 

3,916 

26.958 

32.458 

3.253 

19.430 

4.211 

Skeleton  of  loin 

34.343 

24.734 

3.216 

20.081 

3.905 

Skeleton  of  rib 

3.162 

31.674 

21.579 

3.195 

25.537 

3.726 

Horns 

468 

53.053 

0.655 

5.345 

13.848 

2.689 

Hoofs  and  dewclaws 

869 

58.119 

0.599 

6.018 

1.249 

0.061 

Teeth 

304 

46.836 

0.937 

1.458 

40.727 

7.826 

Studies  In  Animal  Nutrition — III 


51 


Table  22. — Steer  547.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

8,711 

80.850 

2.942 

0.717 

0.029 

Circulatory  system 

L624 

58.440 

27.812 

1.966 

0.682 

0.124 

Respiratory  system 

1,868 

78.906 

2.817 

2.547 

1.149 

0.195 

Brain  and  spinal  cord 

459 

77.158 

8.963 

1.537 

1.460 

0.367 

Digestive  and  excretory  system  (partial) 

11,978 

74.163 

10.929 

2.138 

1.042 

0.192 

Offal  fat 

3,879 

18.056 

78.881 

0.660 

0.257 

0.053 

Liver 

2,851 

71.964 

2.581 

2.956 

1.364 

0.366 

Spleen 

391 

77.086 

2.212 

3.020 

1.345 

0.299 

Pancreas 

200 

71.405 

10.115 

2.426 

1.455 

0.376 

Kidneys 

450 

73 . 255 

8.553 

2.560 

1.144 

0.240 

Hair  and  hide 

14,618 

64.546 

1.829 

5.330 

1.295 

0.072 

Head,  tail,  shin  and  shank,  lean  and  fat 

8.590 

68.763 

12.261 

2.818 

0.897 

0.173 

Flank  and  plate,  lean  and  fat 

14,226 

55.003 

27.523 

2.502 

0.793 

0.147 

Rump,  lean  and  fat 

2.256 

54.409 

27.572 

2.464 

0.818 

0.157 

Chuck  and  neck,  lean  and  fat 

23,636 

67.588 

12.501 

2.772 

0.918 

0.167 

Round,  lean 

17.092 

74.440 

3.584 

3.223 

1.079 

0.210 

Round,  fat 

2,150 

29.872 

60.663 

1.343 

0.397 

0.068 

Loin,  lean 

13,576 

71.512 

6.124 

3.182 

1.030 

0.199 

Loin,  fat 

3,972 

21.083 

72.247 

0.987 

0.300 

0.057 

Rib,  lean 

6.560 

69.801 

9.433 

3.043 

0.979 

0.180 

Rib, fat 

1,102 

25.085 

66.287 

1.438 

0.503 

0.075 

Kidney,  fat 

1,630 

7.887 

90.103 

0.312 

0.136 

0.015 

Skeleton  of  feet,  head,  tail,  shin  and  shank.. . 

11,996 

44.993 

14.086 

3.341 

18.989 

3.501 

Skeleton  of  flank  and  plate 

1,958 

53.339 

13.420 

3.110 

11.771 

2.168 

Skeleton  of  rump 

760 

37.716 

15.923 

3.294 

23.682 

4.444 

Skeleton  of  chuck  and  neck 

5,120 

43.990 

13.982 

3.220 

20.226 

3.645 

Skeleton  of  round 

2,488 

36.276 

28.609 

2.525 

17.314 

3.258 

Skeleton  of  loin 

3.132 

39.384 

20.100 

2.909 

20.155 

3.841 

Skeleton  of  rib 

2,172 

41.532 

16.945 

3.321 

19.474 

3.632 

Horns,  hoofs  and  dewclaws 

737 

53.209 

1.331 

7.310 

2.465 

0.220 

Teeth 

310 

32.500 

0.680 

2.200 

51.200 

9.550 

Table  23. — Steer  548.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

4,603 

753 

81.198 

3.100 

0.715 

0.021 

Circulatory  system 

70 . 180 

12.580 

2.606 

0.843 

0.157 

Respiratory  system 

1,084 

76.883 

3.131 

2.908 

1.106 

0.202 

Brain  and  spinal  cord 

526 

74 . 634 

11.062 

1.637 

1.434 

0.350 

Digestive  and  excretory  system,  partial 

Offal  fat 

5,357 

845 

78.972 

52.573 

4.268 

36.778 

2.386 

1.461 

1.066 

0.720 

0.179 

0.122 

Heart  and  neck  sweetbreads 

168 

77.090 

6.097 

2.816 

1.577 

0.392 

Liver 

1,104 

70.817 

1.835 

3.286 

1.381 

0.336 

Spleen 

215 

77.264 

1.368 

3.131 

1.342 

0.281 

Pancreas 

85 

75.456 

6.206 

2.812 

1.394 

0.331 

Kidneys 

353 

78.549 

3.384 

2.489 

1.237 

0.247 

Hair  and  hide 

8,358 

65.408 

0.933 

5.346 

1.349 

0.078 

Head  and  tail,  lean  and  fat 

967 

71.688 

8.999 

2.675 

0.953 

0.175 

Shin  and  shank,  lean  and  fat 

2,462 

74.845 

4.448 

3.267 

0.998 

0.175 

Flank  and  plate,  lean  and  fat 

4,802 

70.383 

7.627 

3.320 

0.968 

0.174 

Rump,  lean  and  fat 

1,402 

11,136 

9,208 

520 

72.577 

5.800 

3.353 

1.106 

0.204 

Chuck  and  neck,  lean  and  fat 

75.625 

3.506 

3.100 

1.053 

0.187 

Round,  lean 

75.514 

1.680 

3.363 

1.118 

0.214 

Round,  fat 

53.819 

28.733 

2.739 

0.692 

0.086 

Loin,  lean 

5,668 

384 

75.325 

2.020 

3.292 

1.128 

0.209 

Loin,  fat 

40 . 939 

45.571 

1.971 

0.586 

0.097 

Rib,  lean  and  fat 

3,180 

75.848 

2.738 

3.133 

1.093 

0.195 

Kidney,  fat 

284 

27.638 

64.220 

1.384 

0.490 

0.060 

Skeleton  of  feet 

2.496 

2,989 

94 

45 . 620 

13.826 

3.344 

18.414 

3.215 

Skeleton  of  head 

56.949 

6.785 

2.771 

17.108 

3.133 

Skeleton  of  tail 

56.745 

10.923 

3.378 

10.652 

1.934 

Skeleton  of  shin 

1,584 

1,712 

1,514 

570 

38.976 

19.708 

3.222 

19.487 

3.568 

Skeleton  of  shank 

35.963 

21.195 

3.613 

18  669 

3.466 

Skeleton  of  flank  and  plate 

57.844 

10.775 

3.196 

8.981 

1.546 

Skeleton  of  rump 

43.892 

15.924 

3.252 

18.272 

3.323 

Skeleton  of  chuck  and  neck 

3.630 

47 . 340 

13  202 

3.109 

17.177 

3.208 

Skeleton  of  round 

1,898 
1 ,652 

40.419 

26.174 

2.620 

16.404 

3.017 

Skeleton  of  loin 

43.992 

18.943 

3.128 

17  030 

3.230 

Skeleton  of  rib 

1,614 

46  672 

14.692 

3.420 

16.464 

3.005 

Horns,  hoofs  and  dewclawB 

435 

55.591 

1 . 144 

6.927 

2.606 

0.264 

Teeth 

264 

42.122 

0.624 

2.907 

42.160 

7.937 

52 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  24. — Steer  550.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

7,080 

81.361 

2.867 

0 608 

0 031 

Circulatory  system 

1,163 

55.997 

29.913 

1.989 

0.660 

0.133 

Respiratory  system 

1,460 

77.875 

3.987 

2.478 

0.997 

0.199 

Brain  and  spinal  cord 

443 

75.197 

10.097 

1.581 

1.440 

0.351 

Digestive  and  excretory  system  (partial) .... 

9,156 

72.665 

13.019 

1.955 

1.176 

0.185 

Offal  fat 

2.610 

20.877 

74.687 

0.587 

0.323 

0.079 

Liver 

1,992 

70.203 

3.137 

2.994 

1.324 

0.312 

Spleen 

291 

76.774 

2.659 

2.941 

1.480 

0.325 

Pancreas 

200 

66.906 

15.542 

2.442 

1.338 

0.320 

Kidneys 

379 

75.357 

6.259 

2.503 

1.173 

0.252 

Hair  and  hide 

10,440 

64.935 

1.217 

5.320 

1.307 

0.084 

Head,  tail,  shin  and  shank,  lean  and  fat 

5,274 

71.146 

9.321 

2.828 

0.946 

0.174 

Plank  and  plate,  lean  and  fat 

7,896 

61.653 

19.406 

2.857 

0.888 

0.164 

Rump,  lean  and  fat 

1,662 

59.888 

20.746 

2.688 

0.918 

0.178 

Chuck  and  neck,  lean  and  fat 

15,814 

67.239 

14.099 

2.685 

0.950 

0.180 

Round,  lean 

11  720 

75.505 

2.653 

3.418 

1.063 

0.213 

Round,  fat 

944 

32.441 

57.921 

1.439 

0.416 

0.069 

Loin,  lean 

9.072 

73.955 

4.503 

3.097 

1.059 

0.231 

Loin,  fat 

2,134 

21.819 

71.450 

0.918 

0.326 

0.065 

Rib,  lean  and  fat 

4,408 

69.082 

11.035 

2.924 

0.996 

0.187 

Kidney, fat 

756 

10.430 

86.878 

0.455 

0.229 

0.057 

Skeleton  of  feet,  head,  tail,  shin  and  shank.. . 

9,839 

45.940 

15.646 

2.949 

18.669 

3.461 

Skeleton  of  flank  and  plate 

1,540 

52.599 

12.791 

3.242 

12.161 

2.167 

Skeleton  of  rump 

700 

40.804 

16.240 

3.381 

19.007 

3.635 

Skeleton  of  chuck  and  neck 

4,956 

44.877 

16.362 

3.097 

18.062 

3.294 

Skeleton  of  round 

2,100 

36.994 

28.692 

2.629 

16.908 

3.207 

Skeleton  of  loin 

2,464 

38.480 

24.756 

2.896 

17.775 

3.340 

Skeleton  of  rib 

1,740 

41.987 

17.627 

3.296 

18.509 

3.404 

Horns,  hoofs  and  dewclaws 

549 

53.209 

1.331 

7.310 

2.465 

0.220 

Teeth 

228 

32.500 

0.680 

2.200 

51.200 

9.550 

Table  25. — Steer  552.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

5,219 

79.774 

3.010 

0.854 

0.028 

Circulatory  system 

1,056 

64.067 

20.784 

2.167 

0.788 

0.130 

Respiratory  system 

1.096 

77.900 

2.614 

2.724 

1.206 

0.205 

Brain  and  spinal  cord 

466 

74.074 

11.478 

1.564 

1.369 

0.335 

Digestive  and  excretory  system  (partial) 

5,936 

75.009 

9.539 

2.159 

1.058 

0.174 

Offal  fat 

1,784 

28.924 

67.756 

0.924 

0.380 

0.069 

Heart  and  neck  sweetbreads 

265 

62.652 

23.511 

2.325 

1.439 

0.358 

Liver 

1,181 

69.513 

2.065 

3.127 

1.333 

0.337 

Spleen 

284 

77.867 

1.297 

3.100 

1.340 

0.279 

Pancreas 

101 

73.221 

9.258 

2.422 

1.292 

0.301 

Kidneys 

316 

72.871 

12.099 

2.305 

1.128 

0.221 

Hair  and  hide 

10,532 

66.823 

1.534 

4.828 

1.405 

0.070 

Head  and  tail,  lean  and  fat 

1,209 

68.009 

14.483 

2.476 

0.932 

0.155 

Shin  and  shank,  lean  and  fat 

2,976 

74.470 

4.636 

3.211 

0.976 

0.172 

Plank  and  plate,  lean  and  fat 

6,204 

63.328 

17.537 

2.866 

0.882 

0.145 

Rump,  lean  and  fat 

1,326 

65.584 

14.803 

2.886 

0.976 

0.175 

Chuck  and  neck,  lean  and  fat 

12,684 

72.915 

7.088 

2.995 

0.994 

0.178 

Round,  lean 

9,830 

76.090 

1.957 

3.255 

1.129 

0.207 

Round,  fat 

908 

44.576 

41.667 

2.111 

0.589 

0.067 

Loin,  lean 

7,034 

74.559 

3.588 

3.131 

1.101 

0.194 

Loin,  fat 

1,042 

22.439 

70.990 

1.179 

0.395 

0.061 

Rib,  lean  and  fat 

4,104 

72.405 

6.435 

3.233 

0.969 

0.180 

Kidney, fat 

500 

12.924 

84.558 

0.535 

0.281 

0.034 

Skeleton  of  feet 

2,861 

43.452 

12.823 

3.478 

19.568 

3.480 

Skeleton  of  head 

3,052 

54.140 

6.234 

2.916 

18.811 

3.453 

Skeleton  of  tail 

131 

53.865 

13.364 

2.887 

12.790 

2.320 

Skeleton  of  shin 

1,550 

39.817 

17.664 

3.553 

18.794 

3.332 

Skeleton  of  shank 

1,742 

32.535 

20.841 

3.086 

24.917 

4.583 

Skeleton  of  flank  and  plate 

1,516 

54.273 

12.715 

3.371 

10.652 

1.911 

Skeleton  of  rump 

640 

39.791 

17.248 

3.300 

20.750 

3.671 

Skeleton  of  chuck  and  neck 

3,908 

44.753 

15.051 

3.224 

18.379 

3.344 

Skeleton  of  round 

1.626 

33.311 

29.673 

2.565 

19.808 

3.484 

Skeleton  of  loin 

2.192 

41.054 

20.912 

2.874 

18.374 

3.457 

Skeleton  of  rib 

2.084 

43.800 

16.187 

3.281 

17.515 

3.306 

Horns,  hoofs  and  dewclaws 

550 

57.684 

1.828 

6.410 

2.692 

0.261 

Teeth 

278 

40.922 

0.772 

2.035 

43.221 

8.211 

Studies  In  Animal  Nutrition — III 


53 


Table  26. — Steer  554.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

4,197 

591 

81.073 

2.890 

0.801 

0.031 

72.575 

10.404 

2.535 

0.908 

0.169 

907 

79.018 

2.438 

2.691 

1.209 

0.227 

395 

76.826 

8.780 

1.774 

1.581 

0.350 

Digestive  and  excretory  system  (partial) .... 

4,216 

644 

77.716 

41.234 

7.707 

50.872 

2.132 

1.202 

1.098 

0.634 

0.189 

0.099 

336 

80.688 

2.135 

2.651 

2.125 

0.474 

1,166 

202 

70.740 

2.587 

2.813 

1.688 

0 343 

78.308 

1.436 

2.961 

1.484 

0 312 

70.340 

13.270 

2.357 

1.501 

0 280 

530 

81.509 

2.392 

2.221 

1 . 152 

0 215 

Hair  and  hide 

7,400 

883 

66.018 

1.615 

4.976 

1.442 

0 096 

Head  and  tail, lean  and  fat 

72.777 

8.344 

2.812 

1.253 

0 191 

Shin  and  shank,  lean  and  fat 

2,304 

74.453 

3.945 

3.321 

1.126 

0.176 

Flank  and  plate,  lean  and  fat 

4,232 

70.739 

8.896 

2.995 

0.980 

0.170 

Rump,  lean  and  fat 

1,052 

70.442 

9.091 

3.057 

1.166 

0.178 

Chuck  and  neck,  lean  and  fat 

10.530 

7,918 

520 

74.853 

4.416 

3.102 

1.245 

0.189 

Round,  lean 

75.950 

2.163 

3.286 

1.271 

0.215 

Round  fat 

51.440 

32.679 

2.668 

0.762 

0 083 

Loin  lean  

5,812 

428 

74.847 

3.700 

3.199 

1.150 

0.207 

Loin  fat 

29.732 

57.870 

1 394 

0 536 

0 079 

Rib,  lean  and  fat 

Kidney,  fat 

2,914 

240 

74.793 

18.564 

3.459 

75.417 

3.261 

0.727 

1.138 

0.344 

0.198 

0.057 

Skeleton  of  feet 

2,403 

46.494 

13.980 

3.702 

17.687 

3.284 

Skeleton  of  head 

2.229 

125 

59.707 

3.158 

3.111 

16.850 

3 152 

Skeleton  of  tail 

56.788 

11.330 

3.423 

11  074 

2 065 

Skeleton  of  shin 

1.464 

40  004 

18.416 

3.221 

20.982 

3.982 

Skeleton  of  shank 

1.918 

40.266 

19.360 

3.248 

17.454 

3.124 

Skeleton  of  flank  and  plate 

1.278 

60  681 

9.953 

3 288 

7.753 

1.374 

Skeleton  of  rump 

732 

48.799 

12.664 

3.475 

16.288 

2.950 

Skeleton  of  chuck  and  neck 

3,732 

51.461 

10.035 

3.153 

16.389 

2.945 

Skeleton  of  round 

1,702 

1.868 

1.354 

338 

40 . 198 

24.701 

2.669 

15.953 

2 707 

Skeleton  of  loin 

50 . 157 

13.850 

2.994 

14.999 

2 690 

Skeleton  of  rib 

50.983 

12.314 

3.175 

15.340 

2 660 

Horns,  hoofs  and  dewclaws 

55.271 

1.451 

7.670 

2.390 

0.229 

Teeth. 

225 

46.326 

0.098 

2.946 

38.711 

6.360 

Table  27. — Steer  555.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

4,529 

554 

80.075 

3 124 

0.738 

0.035 

Circulatory  system 

70 . 982 

12  062 

2.574 

0.875 

0.159 

Respiratory  system 

937 

79.445 

2.779 

2.693 

1.115 

0.208 

Brain  and  spinal  cord 

353 

76.371 

9.945 

1.705 

1.535 

0.354 

Digestive  and  excretory  system  (partial) .... 
Offal  fat 

4,534 

462 

78.376 
58  692 

6.357 

31.373 

2.319 

1.720 

1.091 

0.779 

0.189 

0.119 

Heart  and  neck  sweetbreads 

159 

75  984 

6.375 

2.732 

1.347 

0.274 

Liver 

1,240 

167 

74  635 

2.536 

2.933 

1 493 

0.349 

Spleen 

77.376 

1.363 

3.129 

1.308 

0.276 

Pancreas 

106 

76.261 

5 510 

2 622 

1.458 

0.293 

Kidneys 

439 

81.997 

2.743 

2.220 

1.181 

0.223 

Hair  and  hide 

6,580 

956 

69.051 

0 . 755 

5.200 

1.476 

0.089 

Head  and  tail,  lean  and  fat 

72.679 

9.234 

2.630 

1.055 

0.162 

Shin  and  shank,  lean  and  fat 

2,688 

4,068 

876 

77.046 

2.331 

3.187 

1.044 

0.181 

Flank  and  plate,  lean  and  fat 

75.514 

3.424 

3.179 

1.038 

0.175 

Rump,  lean  and  fat 

71.979 

7.274 

2.915 

1.070 

0.187 

Chuck  and  neck,  lean  and  fat 

9,402 

7,126 

376 

77  450 

2 143 

3.130 

1 041 

0.185 

Round,  lean 

77  949 

1 347 

3.059 

1.194 

0.207 

Round,  fat 

58  987 

23.814 

2.599 

0.747 

0.090 

Loin,  lean 

4,618 

274 

77.939 

1.726 

3.105 

1.200 

0.208 

Loin,  fat 

41  153 

42.967 

2.179 

0.822 

0.100 

Rib,  lean  and  fat 

2,512 

130 

77  (11  1 

2 . 536 

3.101 

1.174 

0.189 

Kidney, fat 

32 . 970 

57.414 

1.209 

0.549 

0 091 

Skeleton  of  feet 

2,157 

51  458 

9.040 

4.041 

16  248 

2 781 

Skeleton  of  head 

1,978 

74 

62.439 

3.214 

2.859 

14.812 

2.570 

Skeleton  of  tail 

63 . 368 

6.593 

3.646 

9.138 

1.523 

Skeleton  of  shin 

1,454 

50.339 

1 1 063 

3.234 

17.828 

3.190 

Skeleton  of  shank 

1,704 

49.701 

13.806 

2.882 

14.863 

2.656 

54 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  27. — Steer  555.  Analysis  of  Samples — Continued. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Skeleton  of  flank  and  plate 

1,236 

506 

65.173 

5.690 

3.369 

8.036 

1.361 

Skeleton  of  rump 

57 . 102 

5.911 

3.496 

16.138 

2.764 

Skeleton  of  chuck  and  neck 

3,216 

1,652 

59.309 

5.153 

3.142 

13.912 

2.463 

Skeleton  of  round 

53.566 

13.254 

2.793 

14.801 

2.582 

Skeleton  of  loin 

1,264 

1.256 

59.306 

7.164 

3.098 

13.007 

2.313 

Skeleton  of  rib 

58.491 

6.345 

3.269 

13.194 

2.272 

Horns,  hoofs  and  dewclaws 

298 

48.852 

0.914 

7.593 

2.662 

0.133 

Teeth 

190 

42.647 

0.330 

2.870 

42.862 

7.297 

Table  28. — Steer  556.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood  

6,124 

81.472 

2.778 

0.784 

0.035 

Circulatory  system 

993 

62.355 

23.955 

2.146 

0.856 

0.148 

Respiratory  system 

1,272 

77.882 

3.420 

2.698 

1.400 

0.216 

Brain  and  spinal  cord 

398 

67.120 

19.452 

1.624 

1.616 

0.349 

Digestive  and  excretory  system  (partial) .... 

6,003 

74.523 

10.512 

2.203 

1.304 

0.241 

Offal  fat 

1,402 

32.618 

61.309 

0.961 

0.451 

0.063 

Heart  and  neck  sweetbreads 

328 

76.405 

6.578 

2.590 

2.052 

0.452 

Liver 

1,760 

71.112 

3.500 

3.060 

2.295 

0.366 

Spleen 

300 

77.717 

2.323 

2.983 

1.480 

0.272 

Pancreas 

96 

72.244 

10.649 

2.415 

1.503 

0.302 

Kidneys 

338 

75.331 

7.087 

2.522 

1.345 

0.254 

Hair  and  hide 

10,314 

66.284 

2.153 

5.157 

1.377 

0.088 

Head  and  tail,  lean  and  fat 

914 

68.746 

13.192 

2.739 

0.897 

0.165 

Shin  and  shank,  lean  and  fat 

2,808 

73.731 

4.508 

3.271 

0.966 

0.185 

Flank  and  plate,  lean  and  fat 

6,174 

67.267 

13.359 

3.143 

0.935 

0.168 

Rump,  lean  and  fat 

1,244 

69.336 

10.388 

3.015 

1.036 

0.191 

Chuck  and  neck,  lean  and  fat 

12,406 

72.316 

7.615 

3.016 

1.246 

0.180 

Round,  lean 

9,472 

75.179 

3.195 

3.289 

1.251 

0.210 

Round,  fat 

686 

44.562 

43.541 

2.194 

0.577 

0.069 

Loin,  lean 

6,938 

74.185 

3.780 

3.270 

1.189 

0.206 

Loin,  fat 

820 

26.114 

66.035 

1.399 

0.542 

0.064 

Rib,  lean  and  fat 

3,300 

71.440 

7.223 

3.184 

1.153 

0.184 

Kidney,  fat 

420 

15.362 

80.619 

0.587 

0.355 

0.037 

Skeleton  of  feet 

2,589 

46.442 

14.176 

3.729 

17.758 

3.279 

Skeleton  of  head 

2,610 

56.628 

7.187 

2.990 

17.629 

3.271 

Skeleton  of  tail 

92 

56.900 

9.965 

3.538 

13.365 

2.393 

Skeleton  of  shin 

1,702 

40.276 

19.800 

3.280 

19.776 

3.695 

Skeleton  of  shank 

1,952 

37.454 

22.610 

3.389 

19.971 

3.787 

Skeleton  of  flank  and  plate 

1,578 

57.659 

11.660 

3.313 

10.266 

1.862 

Skeleton  of  rump 

608 

47.189 

13.166 

3.537 

18.150 

3.408 

Skeleton  of  chuck  and  neck 

3,734 

47.104 

12.535 

3.488 

18.545 

3.545 

Skeleton  of  round 

1,974 

38.147 

25.725 

2.665 

16.702 

3.212 

Skeleton  of  loin 

2,268 

46.157 

15.957 

3.158 

16.929 

3.309 

Skeleton  of  rib 

1,646 

46.368 

12.349 

3.310 

18.157 

3.720 

Horns,  hoofs  and  dewclaws 

390 

48.189 

1.236 

8.303 

1.870 

0.156 

Teeth 

218 

41.753 

0.042 

2.938 

43.927 

6.910 

Studies  In  Animal  Nutrition — III 


55 


Table  29. — Steer  557.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

8,952 

79.941 

3.136 

0.842 

0.027 

Circulatory  system 

2,342 

50.516 

38.557 

1.650 

0.615 

0.105 

Respiratory  system 

1.972 

77.053 

5.258 

2.598 

1.119 

0.203 

Brain  and  spinal  cord 

551 

72.578 

12.435 

1.790 

1.410 

0.338 

Digestive  and  excretory  system  (partial) .... 

9.981 

73.851 

11.446 

2.007 

1.051 

0.176 

Offal  fat 

6.757 

13.945 

83.471 

0.435 

0.229 

0.029 

Heart  and  neck  sweetbreads 

623 

67.787 

16.888 

2.328 

1.653 

0.357 

I.iver 

3,003 

69.463 

1.882 

3.084 

1 344 

0.332 

Spleen 

485 

76.934 

3.341 

2.888 

1.372 

0.281 

Pancreas 

225 

64.844 

17.153 

2.276 

1.068 

0.251 

Kidneys 

649 

76.753 

5.885 

2.376 

1.151 

0.218 

Hair  and  hide 

14,100 

63.124 

5.012 

4.988 

1.171 

0.071 

Head  and  tail,  lean  and  fat 

1.915 

60.473 

24 . 133 

2.352 

0.848 

0.144 

Shin  and  shank,  lean  and  fat 

4,196 

68.205 

11.827 

2.933 

0.927 

0.153 

Flank  and  plate,  lean  and  fat 

15,294 

50.434 

34.335 

2.334 

0.778 

0.133 

Rump,  lean  and  fat 

2,582 

51.177 

32.845 

2 317 

0.797 

0.142 

Chuck  and  neck,  lean  and  fat 

22.146 

63.903 

17.574 

2.702 

0.957 

0.183 

Round,  lean 

14,960 

73.824 

4.610 

3.073 

1.011 

0.194 

Round,  fat 

2,532 

27.746 

63.644 

1.019 

0.349 

0.048 

Loin,  lean 

11.714 

71 . 155 

7.826 

2.966 

1.098 

0.192 

Loin,  fat 

4.472 

15.907 

79.899 

0.729 

0.248 

0.038 

Rib,  lean 

6.372 

67.831 

11  987 

2.905 

0.958 

0.170 

Rib,  fat 

1,616 

19.571 

73.914 

0.959 

0.360 

0.059 

Kidney,  fat 

3.228 

8.267 

89.768 

0.367 

0.159 

0.024 

Skeleton  of  feet 

3,558 

44.384 

11.995 

3.601 

18.856 

3.488 

Skeleton  of  head 

3,969 

53.816 

6.263 

2.751 

18.265 

3.294 

Skeleton  of  tail 

193 

53.009 

16.791 

3.163 

10.458 

1.954 

Skeleton  of  shin 

2,246 

36.792 

18.013 

3.351 

21.238 

3.961 

Skeleton  of  shank 

2,610 

36.883 

19.539 

3.405 

20.052 

3.625 

Skeleton  of  flank  and  plate 

2,498 

56.391 

12.656 

3.152 

9.709 

1.721 

Skeleton  of  rump 

960 

42.749 

13.401 

3.441 

19.159 

3.710 

Skeleton  of  chuck  and  neck 

5,638 

42.943 

12.074 

3.394 

20.847 

3.764 

Skeleton  of  round 

2,686 

32.505 

29.395 

2.573 

19.164 

3.548 

Skeleton  of  loin 

2,478 

38.626 

20.143 

2.959 

19.381 

3.655 

Skeleton  of  rib 

2,572 

40.350 

16.831 

3.239 

19.671 

3.611 

Horns,  hoofs  and  dewclaws 

695 

53.666 

1.415 

6.956 

2.569 

0.279 

Teeth 

261 

39.979 

0.283 

2.874 

46.115 

8.632 

Table  30. — Steer  558.  Analysis  of  Samples. 


Description  of  sample 

Weight  in 
animal, 
grams 

Moisture 

% 

Crude  fat 

% 

Nitrogen 

% 

Ash 

% 

Phosphorus 

% 

Blood 

4,666 

83 . 665 

2.509 

0.692 

0.039 

Circulatory  system 

971 

66.245 

19.036 

2.219 

0.726 

0.143 

Respiratory  system 

1,111 

79.632 

1.472 

2.685 

1.054 

0.231 

Brain  and  spinal  cord 

520 

75.596 

10.002 

1.596 

1.507 

0.374 

Digestive  and  excretory  system  (partial) .... 

6,510 

77.962 

6.366 

2.241 

0.992 

0.188 

Offal  fat 

829 

47.598 

43.206 

1.386 

0.505 

0.106 

Liver 

1,352 

71.470 

2.100 

3.096 

1.349 

0.352 

Spleen 

193 

75.805 

1.099 

3.298 

1.537 

0.278 

Pancreas 

153 

76.282 

5.186 

2.553 

1.452 

0.374 

Kidneys 

318 

76.794 

4.347 

2.595 

1.193 

0.247 

Hair  and  hide 

8,138 

64.713 

1.078 

5.287 

1 . 130 

0.083 

Head,  tail,  shin  and  shank,  lean  and  fat 

4,324 

74.380 

5.509 

2.948 

0.943 

0.163 

Flank  and  plate,  lean  and  fat 

4,192 

70.650 

7.311 

3.229 

0.923 

0.172 

Rump,  lean  and  fat 

1,030 

69.840 

9.636 

3.089 

1.011 

0.196 

Chuck  and  neck,  lean  and  fat 

11,682 

75.244 

4.121 

3.041 

0.927 

0.187 

Round,  lean 

9,664 

77.152 

1.113 

3.116 

1.049 

0.213 

Round,  fat 

644 

44.994 

41.335 

2.308 

0.611 

0.095 

Loin,  lean 

5,952 

74  950 

2.652 

3.182 

1.025 

0 210 

Loin,  fat 

600 

30.204 

59.420 

1.549 

0.446 

0.080 

Rib,  lean  and  fat 

3,242 

74.678 

3.490 

3.157 

0.990 

0.197 

Kidney,  fat 

220 

18.408 

75.158 

0.936 

0.283 

0.055 

Skeleton  of  feet,  head,  tail,  shin  and  shank  . . 

9,785 

45.737 

16  814 

3.097 

17.541 

3.252 

Skeleton  of  flank  and  plate 

1,092 

54.633 

1 - 519 

3.306 

10.416 

1.885 

Skeleton  of  rump 

678 

42. 549 

18.378 

3.113 

18.671 

3.383 

Skeleton  of  cnuck  and  neck 

4,056 

43  212 

18  090 

3.089 

17.848 

3.240 

Skeleton  of  round 

2,094 

34.269 

33.809 

2.285 

16.102 

2.982 

Skeleton  of  loin 

2,138 

39  771 

25.039 

2.529 

17  837 

3.281 

Skeleton  of  rib 

1,516 

46.885 

16  047 

3.348 

15.188 

2.747 

Homs,  hoofs  and  dewclaws 

443 

53.209 

1.331 

7.310 

2.465 

0.220 

Teetn 

274 

32.500 

0.680 

2.200 

51.200 

0.550 

56 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  31. — Steer  500.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Pnospnorus 

Blood 

21,269 

16811.2 

40.8 

679.1 

167.8 

4.68 

Circulatory  system 

1,562 

756.8 

587.9 

29.2 

8.5 

0.95 

Lean  neart 

1,284 

995.7 

45.7 

35.0 

14.2 

2.67 

Respiratory  system 

3,747 

2866.5 

101.2 

106.4 

42.9 

6.45 

Fat  from  thoracic  cavity 

568 

96.8 

450.9 

2.8 

1.4 

0.14 

Brain  and  spinal  cord 

832 

522.2 

180.7 

13.9 

14.5 

3.09 

Digestive  and  excretory  system  (partial) 

19,275 

14385.9 

1921.9 

424.6 

160.8 

22.75 

Offal  fat 

12,940 

1747.0 

10812.2 

54.6 

23.8 

2.59 

Heart  and  neck  sweetbreads 

538 

286.2 

182.6 

9.9 

5.6 

1.21 

Liver 

4.634 

3233.9 

134.5 

150.3 

73.2 

14.97 

Gall 

241 

221.4 

0.5 

0.5 

3.0 

0.07 

Spleen 

1,054 

783.9 

56.4 

31.4 

12.6 

2.31 

Pancreas 

625 

369.6 

156.8 

13.4 

7.5 

1.60 

Kidneys 

1,019 

785.6 

49.0 

24.7 

11.4 

2.11 

Tongue,  marketable 

1,619 

1123.7 

192.2 

43.8 

14.8 

2.49 

Hair  and  hide 

35,938 

21320.9 

474.0 

2256.9 

385.3 

15.81 

Head  and  tail,  lean  and  fat 

3,784 

2410.9 

603.9 

119.4 

33.4 

5.07 

Shin  and  shank,  lean  and  fat 

12,496 

8854.9 

823.6 

420.2 

123.6 

20.49 

Flank  and  plate,  lean  and  fat 

36,410 

19948.3 

10067.7 

978.3 

315.3 

50.61 

Rump,  lean  and  fat 

7,058 

3892.4 

1950.8 

178.4 

57.8 

10.23 

Chuck  and  neck,  lean  and  fat 

58,918 

39825.0 

6993.0 

1995.6 

537.3 

93.09 

Round,  lean 

39,898 

29536.9 

1390.5 

1246.0 

403.4 

76.21 

Round,  fat 

4,936 

1370.6 

3032.8 

78.5 

18.6 

2.52 

Loin,  lean 

29,692 

20864.3 

2296.4 

924.3 

299.9 

54.93 

Loin,  fat 

6,830 

1124.5 

5225.5 

40.8 

16.7 

2.60 

Rib,  lean 

13,602 

9132.0 

1676.2 

434.7 

126.4 

23.12 

Rib,  fat 

1,804 

367.4 

1282.4 

23.3 

6.7 

1.08 

Kidney,  fat 

2,432 

170.9 

2195.5 

10.0 

3.5 

0.44 

Skeleton  of  feet 

6,838 

2708.1 

788.3 

247.0 

1707.5 

309.69 

Skeleton  of  head 

8,953 

4296.2 

1216.2 

312.2 

1599.2 

307.45 

Skeleton  of  tail 

386 

151.7 

92.7 

10.2 

61.4 

10.75 

Skeleton  of  shin 

5,610 

1485.9 

1210.9 

207.6 

1651.6 

292.34 

Ske*eton  of  snank 

5 750 

1808.8 

1160.8 

198.6 

1448.0 

254.67 

Skeieton  of  flank  and  plate 

6,350 

2605.5 

1143.5 

204.7 

1177.1 

203.20 

Skeleton  of  rump 

2,988 

727.3 

914.6 

91.6 

749.8 

132.37 

Skeleton  of  chuck  and  neck 

14,450 

4302.5 

3253.7 

442.2 

3746.3 

661.09 

Skeleton  of  round,  (excl.  marrow) 

6,438 

2095.7 

1789.3 

166.7 

1356.7 

243.74 

Marrow  from  skeleton  of  round 

680 

64.3 

606.9 

1.0 

1.5 

0.21 

Skeleton  of  loin 

7,772 

1947.4 

2438.5 

228.1 

1865.8 

332.41 

Skeleton  of  rib 

5,192 

1409.4 

1158.2 

165.2 

1449.9 

257.11 

Hoofs  and  dewclaws 

2 095 

1059 . 7 

17.5 

162.2 

54.6 

2.45 

Teeth 

852 

181.7 

9.9 

17.7 

519.8 

98.12 

Table  32. — Steer  501.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

28,710 

1,836 

1,882 

3,838 

22387.2 

50.5 

944.6 

246.0 

7.18 

Circulatory  system 

770.4 

842.5 

35.0 

8.7 

0.90 

Lean  heart 

1460.7 

70.4 

48.4 

18.7 

3.73 

Respiratory  system 

2939.0 

128.5 

110.3 

40.0 

6.60 

Fat  from  thoracic  cavity 

2,459 

757 

463.1 

1884.7 

15.1 

5.8 

0.64 

Brain  and  spinal  cord 

533.0 

100.5 

12.7 

13.9 

2.97 

Digestive  and  excretory  system  (partial) 

Offal  fat 

24,235 

38,625 

784 

17390.1 

2892.2 

3119.0 

35172.3 

519.4 

79.2 

196.1 

39.4 

29.81 

4.64 

Heart  and  neck  sweetbreads 

241.1 

484.2 

7.8 

3.9 

0.84 

Liver 

6,161 

176 

4282.7 

178.6 

199.2 

87.7 

20.58 

Gall 

161.8 

0.1 

0.4 

2.2 

0.06 

Spleen 

1,178 

836 

917.6 

23.0 

32.7 

16.3 

2.82 

Pancreas 

500.5 

205.4 

18.4 

9.6 

2.20 

Kidnevs 

1,037 

805.3 

50.5 

24.3 

10.9 

2.06 

Tongue,  marketable 

2,153 

50,090 

5,224 

17,420 

134,146 

22,226 

110,990 

50,130 

22,284 

45,996 

71,358 

1417.6 

348.8 

56.2 

18.7 

3.38 

Hair  and  hide 

25762.3 

6629.4 

2751.4 

762.4 

24.54 

Head  and  tail,  lean  and  fat 

3156.4 

1083.8 

148.0 

40.1 

6.58 

Shin  and  shank,  lean  and  fat 

10268.9 

3932.2 

471.6 

134.5 

23.17 

Flank  and  plate,  lean  and  fat 

35964.5 

88380.8 

1408.5 

458.8 

76.46 

Rump , lean  and  fat 

6394  2 

13949.0 

253.4 

87.8 

15.33 

Chuck  and  neck,  lean  and  fat 

52940.0 

42652.4 

1692.6 

723.7 

130.97 

Round,  lean 

35041.9 

4690.2 

1549.0 

479.7 

92.74 

Round,  fat 

3754.0 

17434.3 

148.6 

48.6 

5.79 

Loin,  lean 

28773 . 7 

8248.9 

1316.9 

391.4 

74.97 

Loin,  fat 

6444.3 

63281.7 

276.9 

79.9 

12.84 

Rib,  lean 

20,834 

12269.6 

4668.7 

574.8 

164  8 

31.04 

Rib,  fat 

28,322 

19,544 

7,744 

2746.4 

24764.5 

113.6 

38.0 

5.66 

Kidnev,  fat 

1067.5 

18236.7 

37.1 

13.1 

2.15 

Skeleton  of  fppt 

2792.0 

954.5 

273.4 

2023.7 

390.68 

Studies  In  Animal  Nutrition — III 


57 


Table  32. — Steer  501.  Weights  of  Constituents  in  Samples,  Grams — Cont. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

10,462 

304 

4561.8 

1232.0 

343.9 

2487.5 

439.30 

Skeleton  of  tail 

124.3 

58.3 

10.6 

54 . 9 

9.59 

Skeleton  of  shin 

6,170 

2018  3 

875.4 

214.5 

1787.3 

323.12 

Skeleton  of  shank 

7,128 

7,068 

1920.5 

1581.5 

238.7 

1985 . 9 

365.24 

Skeleton  of  flank  and  plate 

2842.3 

1107.0 

234.7 

1328 . 9 

240.52 

Skeleton  of  rump 

3,682 

932  8 

965.2 

116.8 

956.3 

172.61 

Skeleton  of  ctruck  and  neck 

15,778 

4750.1 

2444.8 

576.8 

4478.0 

821.24 

Skeleton  of  round,  (excl.  marrow) 

6,978 

1859.4 

1699.2 

215  3 

190 i .7 

345.27 

Marrow  from  skeleton  of  round 

286 

29.1 

252.8 

0.6 

1.5 

0.24 

Skeleton  of  loin 

8,614 

6,388 

3,354 

2,523 

778 

2214.8 

1940 . 1 

269.4 

2447.3 

436.90 

Skeleton  of  rib 

1816.0 

1173.5 

212.2 

1780.2 

330.96 

Horns 

1240.6 

21.2 

217.0 

762.8 

139.76 

Hoofs  and  dewclaws 

1186.1 

16.6 

213.3 

43.3 

3.61 

Teeth 

172.0 

6.3 

16.1 

465.1 

91.31 

Table  33. — Steer  502.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

• Crude  fat 

Nitrogen 

Asn 

Phosphorus 

Blood 

19,728 

15276.0 

688.9 

142.0 

4.54 

Circulatory  system 

3,446 

2353.1 

500.1 

89.1 

23.4 

4.76 

Respiratory  system 

3,696 

2815.1 

113.0 

104.6 

38.4 

6.14 

Fat  from  thoracic  cavity 

1.271 

273.3 

933.9 

9.6 

3.8 

0.52 

Brain  and  spinal  cord 

800 

552.8 

116.8 

13.7 

14.4 

3.31 

Digestive  and  excretory  system  (partial) 

20,933 

16122.6 

1263.3 

522.3 

264.6 

28.47 

Ofial  fat 

11,377 

1211.8 

9804.7 

53.2 

21.1 

2.62 

Heart  and  neck  sweetbreads 

502 

300.1 

127.6 

11.3 

5.8 

1.36 

Liver 

3,716 

2561.9 

64.2 

122.8 

51.7 

12.41 

Gall 

241 

224.2 

0.1 

0.5 

2.5 

0.07 

Spleen 

921 

711.9 

22.3 

26.1 

16.9 

2.57 

Pancreas 

581 

310.6 

172.1 

12.6 

6.2 

1.39 

Kidneys 

838 

617.3 

55.2 

22.5 

9.3 

1.85 

Hair  and  hide 

39,556 

22665.2 

1190.6 

2600.4 

386.1 

23.34 

Head  and  tail,  lean  and  fat 

4,250 

2745.5 

600.2 

136.0 

35.1 

6.33 

Shin  and  shank,  lean  and  fat 

12,364 

8444.0 

1072.1 

431.0 

109.3 

20.28 

Flank  and  plate,  lean  and  fat 

35,594 

18475.1 

11113.2 

919.4 

247.7 

42.36 

Rump,  lean  and  fat 

8,100 

4538.3 

2059.5 

211.5 

64.6 

11.91 

Chuck  and  neck,  lean  and  fat 

70,744 

46926.6 

9004.3 

2173.3 

651.6 

111.78 

Round,  lean 

44,426 

32054.7 

1799.7 

1468.7 

431.4 

86.63 

Round,  fat 

4,620 

1229.0 

2903.1 

71.6 

14.9 

1.76 

Loin,  lean 

35,104 

24529.6 

2852.5 

1120.9 

342.3 

69.86 

Loin,  fat 

9,144 

1416.4 

7149.0 

95.7 

22.8 

3.29 

Rib,  lean 

18,256 

12114.3 

1846.8 

560.1 

163.0 

29.94 

Rib, fat 

3,338 

689.5 

2336.4 

49.5 

9.4 

1.70 

Kidney,  fat 

2,916 

215.4 

2614.7 

12.3 

7.0 

1.14 

Skeleton  of  feet 

6.982 

2897.3 

852.2 

264.5 

1666.5 

296.04 

Skeleton  of  head 

9,577 

4669.5 

802.4 

305.0 

1934.2 

331.36 

Skeleton  of  tail 

441 

178.3 

99.6 

14.3 

66.5 

12.05 

Skeleton  of  shin 

5,940 

1627.5 

1147.6 

209.8 

1750.3 

311.97 

Skeleton  of  shank 

5.978 

1625.8 

1469.7 

206.5 

1516.2 

269.31 

Skeleton  of  flank  and  plate 

5,590 

2300.6 

726.5 

189.1 

1219.6 

215.83 

Skeleton  of  rump 

2,362 

600.8 

702.3 

71.5 

559.4 

99.96 

Skeleton  of  chuck  and  neck 

15,092 

4643.4 

2755.7 

514.2 

4158.3 

742.22 

Skeleton  of  round  (excl.  marrow) 

6,296 

1667.0 

1664.9 

188.6 

1653.3 

294.72 

Marrow  from  skeleton  of  round 

408 

32.1 

369.8 

0.8 

1.7 

0.30 

Skeleton  of  loin 

7,866 

2068.0 

2085 . 6 

247.7 

2032.5 

359.40 

Skeleton  of  rib 

5,920 

1801.6 

1219.5 

215.9 

1472.0 

260.54 

Horns* 

1,949 

745.9 

11.9 

131.0 

410.3 

72.76 

Hoofs  and  dewclaws 

2,010 

1179.6 

11.5 

133.2 

23.8 

2.41 

Teeth 

1,038 

375.9 

10.6 

16.9 

519.0 

98.43 

•This  sample  was  lost  before  analysis.  The  average  analysis  of  the  horns  of  two  mature  steers  in  the. 

same  group  was  used. 


58 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  34. — Steer  503.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

13,058 

10809.4 

14.4 

353.6 

43.9 

9.92 

Circulatory  system 

1,782 

645.9 

978.6 

22.3 

5.9 

1.07 

Lean  heart 

1.063 

830.6 

39.7 

27.5 

10.1 

2.20 

Respiratory  system 

2,549 

2006.2 

80.4 

67.7 

24.9 

5.28 

Brain  and  spinal  cord 

666 

486.9 

107.3 

11.2 

10.3 

2.62 

Digestive  and  excretory  system  (partial) 

8,761 

6370.1 

964.9 

208.1 

88.7 

18.22 

Offal  fat 

7,385 

1081.3 

6049.8 

38.5 

13.4 

2.58 

Liver 

3,646 

2504.1 

192.0 

109.2 

46.8 

12.18 

Kidneys 

655 

467.4 

77.3 

15.8 

6.8 

1.47 

Stomach 

5,765 

4477.0 

399.2 

127.5 

62.6 

11.93 

Tongue,  marketable 

789 

547.0 

104.7 

20.0 

6.5 

1.34 

Hair  and  hide 

23,008 

15591.4 

591.3 

1102.8 

225.3 

15.18 

Shin,  shank,  head  and  tail,  lean  and  fat 

8,614 

6004.6 

880.6 

275.3 

72.9 

14.46 

Flank  and  plate,  lean  and  fat 

16,290 

9206.8 

4181.0 

446.7 

121.9 

22.17 

Chuck  and  neck,  lean  and  fat 

31,934 

21883.1 

4015.4 

943.3 

278.0 

54.61 

Round  and  rump,  lean 

25,022 

18287.8 

1202.3 

843.2 

256.1 

51.14 

Round  and  rump,  fat 

3,400 

763.3 

2384.4 

41.2 

9.8 

1.62 

Loin,  lean 

17,206 

12215.2 

1454.1 

548.4 

169.1 

32.69 

Loin,  fat 

5,746 

904.9 

4592.0 

44.7 

10.9 

2.01 

Rib,  lean 

8,932 

6223.6 

913.8 

281.7 

82.8 

16.52 

Rib,  fat 

840 

195.7 

562.3 

12.4 

2.7 

0.51 

Kidney,  fat 

2,126 

184.5 

1902.1 

7.2 

2.7 

0.58 

Skeleton 

41,122 

15740.3 

6192.6 

1276.4 

9747.6 

1800.32 

Horns,  hoofs  and  dewclaws 

1,058 

489.7 

21.5 

80.2 

64.1 

6.99 

Teeth 

253 

59.0 

1.3 

5.7 

149.0 

28.54 

Table  35. — Steer  504.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

21,005 

16520.4 

19.5 

700.3 

81.3 

4.62 

Circulatory  system 

1,637 

470.0 

1043.3 

18.2 

6.6 

0.67 

Lean  heart 

1,428 

1094.6 

61.6 

39.6 

14.5 

2.90 

Respiratory  system 

3,628 

2445.6 

540.6 

104.6 

35.3 

6.57 

Brain  and  spinal  cord 

724 

501.2 

109.2 

13.0 

9.8 

2.53 

Digestive  and  excretory  system  (partial) 

17,569 

12041.8 

3169.5 

335.6 

127.6 

25.48 

Offal  fat 

25,105 

3203.4 

21294.1 

83.9 

39.9 

5.77 

Liver 

4,754 

3280.7 

115.8 

155.7 

64.3 

17.02 

Kidneys 

877 

609.3 

112.3 

21.5 

9.3 

1.92 

Stomach 

12.820 

1Q213.7 

1030.7 

218.1 

115.0 

19.36 

Tongue,  marketable 

1,587 

964.1 

372.6 

34.6 

12.1 

2.24 

Hair  and  hide 

41,144 

23982.8 

3320.3 

2272.0 

434.9 

17.69 

Shin,  shank,  head  and  tail,  lean  and  fat 

16,070 

9744.9 

3304.0 

474.2 

129.0 

23.46 

Flank  and  plate,  lean  and  fat 

49,650 

20674.3 

22650.3 

965.7 

284.0 

50.15 

Rump,  lean  and  fat 

10,846 

4416.5 

5077.0 

200.3 

62.3 

11.50 

Chuck  and  neck,  lean  and  fat 

59,808 

34886.0 

14371.9 

1567.0 

453.3 

87.32 

Round,  lean 

37,238 

25884.1 

3429.6 

1194.6 

366.1 

72.24 

Round,  fat 

9,818 

1630.8 

7661.0 

89.0 

23.4 

2.95 

Loin,  lean 

33,676 

22535.9 

4115.2 

1027.5 

318.6 

60.95 

Loin,  fat 

18,340 

2131.1 

15572.5 

97.6 

29.7 

4.59 

Rib,  lean 

18,506 

11710.6 

3242.3 

544.1 

153.8 

30.91 

Rib,  fat 

6,770 

976.2 

5458.7 

56.4 

13.7 

2.10 

Kidney,  fat 

11,400 

547.2 

10709.2 

24.5 

14.4 

1.94 

Skeleton  of  feet,  head,  tail,  shin  and  shank.. . 

23,568 

8496.3 

3214.7 

762.9 

6503.4 

1196.31 

Skeleton  of  flank  and  plate 

4,572 

2916.3 

831.2 

161.1 

718.5 

133.32 

Skeleton  of  rump 

2,428 

624.5 

631.3 

74.6 

664.8 

125.87 

Skeleton  of  chuck  and  neck 

11,176 

3372.9 

1818.3 

393.8 

3259.2 

597.25 

Skeleton  of  round 

4,808 

1052.0 

1320.8 

150.0 

1492.8 

276.70 

Skeleton  of  loin 

5,850 

1745.6 

1292.9 

186.3 

1547.4 

281.50 

Skeleton  of  rib 

5,092 

1657.5 

828.0 

174.0 

1415.1 

259.49 

Horns,  hoofs  and  dewclaws 

2,532 

1759.1 

13.0 

117.0 

61.5 

3.98 

Teeth* 

338 

86.2 

3.6 

6.4 

196.8 

37.57 

♦This  sample  was  lost  before  analysis.  The  average  analysis  of  the  teeth  of  four  steers  of  the  same 
Group  was  used. 


Studies  In  Animal  Nutrition — III 


59 


Table  36. — Steer  505.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

13.810 

11360.1 

48.5 

376.5 

45.4 

3.18 

Circulatory  system 

1.168 

388.1 

689.9 

15.0 

3.2 

0.55 

Lean  heart 

938 

725.8 

47.9 

24.6 

9.1 

1.96 

Respiratory  system 

2,498 

1919.2 

136.8 

67.8 

23.8 

5.05 

Brain  and  spinal  cord 

537 

396.4 

79.1 

9.1 

9.2 

2.21 

Digestive  and  excretory  system  (partial) 

Offal  fat 

8.258 

5936.9 

1058.3 

204.5 

87.0 

18.66 

12,781 

1586.1 

10912.9 

43.5 

15.0 

2.81 

Liver 

3,983 

2712.3 

229.8 

127.7 

52.9 

13.82 

Kidneys 

718 

543.6 

56.3 

17.1 

7.6 

1.62 

Stomach 

8,818 

6812.9 

956.7 

148.2 

76.5 

15.26 

Tongue,  marketable 

1.115 

714.3 

224.8 

26.2 

8.5 

1.74 

Hair  and  hide 

22.884 

14219.7 

1221.1 

1212.2 

159.3 

15.10 

Shin,  shank,  head  and  tail,  lean  and  fat 

9.386 

6047.4 

1445.3 

301.9 

76.8 

15.40 

Flank  and  plate,  lean  and  fat 

24,194 

10577.6 

10345.8 

535.0 

140.3 

28.07 

Chuck  and  neck,  lean  and  fat 

38,344 

23886.0 

7265.8 

1106.6 

318.3 

63.27 

Round  and  rump,  lean 

25  784 

17808.0 

2440.2 

857.6 

251.7 

51.57 

Round  and  rump,  fat 

5,970 

844.2 

4814.2 

27.0 

10.4 

1.91 

Loin,  lean 

19,686 

13423.9 

1965.3 

637.0 

189.6 

38.58 

Loin,  fat 

7.558 

705.4 

6616.8 

40.4 

9.6 

1.83 

Rib,  lean 

11,300 

6979.1 

2168.6 

334.9 

95.0 

19.89 

Rib,  fat 

2.640 

288.1 

2254.2 

17.0 

4.4 

0.87 

Kidney, fat 

5,754 

302.8 

5381.5 

13.6 

4.8 

0.92 

Skeleton 

37,745 

13509.7 

6626.2 

1202.6 

9002.9 

1661.92 

Horns,  hoofs  and  dewclaws 

1,206 

555.9 

12.2 

93.2 

64.4 

7.37 

Teeth 

268 

58.8 

1.7 

6.1 

159.2 

28.67 

Table  37. — Steer  507.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

20,316 

4,278 

15881.8 

697.0 

136.1 

4.47 

Circulatory  system 

2015.7 

1765 . 7 

72  5 

22.9 

4.45 
6.37 
3.11 
41  13 

Respiratory  system 

3,768 

744 

2919.0 

148.0 

108.8 
11  2 

35.9 

Brain  and  spinal  cord 

526.8 

102.2 

13.0 

Digestive  and  excretory  system 

27,417 

11,307 

34,473 

4.038 

19333.1 

3860.0 

574.9 

226.2 

Offal  fat 

1480.8 

9493.6 

46.6 

15.8 

3.28 

16.89 

Hair  and  hide 

20975 . 1 

2141.8 

1960.8 

367.1 

Head  and  tail,  lean  and  fat 

2527.1 

765.2 

114.7 

36.1 

6.51 

Shin  and  shank,  lean  and  fat 

11,860 

36,130 

7,736 

62,530 

8288 . 1 

932.2 

396.5 

109.2 

19.81 

Flank  and  plate,  lean  and  fat 

18765.6 

11692.4 

904.7 

250.4 

44  81 

Rump,  lean  and  fat 

3930.1 

2448.0 

176.0 

55.8 

10  75 

Chuck  and  neck,  lean  and  fat 

41000.3 

9483.3 

1807.7 

539.0 

98  07 

Round,  lean 

39,302 

28583 . 2 

2268.5 

1269.5 

385.6 

75.46 

Round,  fat 

5,378 

1314.7 

3693.1 

58.8 

14.8 

2 10 

Loin,  lean 

29,724 

10,188 

15,788 

2,432 

21013.7 

2406.5 

929.8 

288.9 

54  99 

Loin,  fat 

1815.7 

7812.6 

84.2 

22.7 

3 97 

Rib,  lean 

10647.1 

1932.3 

473.5 

147  9 

27.47 

Rib,  fat 

426.9 

1858.9 

23.0 

6.2 

1.02 

Kidney,  fat 

4,376 

15,275 

296.9 

3995.3 

12.4 

6.4 

1.09 

Skeleton  of  feet,  head  and  tail 

6538.3 

1798.8 

527.9 

3651.6 

589.31 

Skeleton  of  shin  and  shank 

10,350 

6,278 

2,536 

2479.9 

2003.5 

377.4 

3489.7 

479.62 

Skeleton  of  flank  and  plate 

2775.9 

846.6 

212.2 

1159.7 

178.99 

Skeleton  of  rump 

635  0 

668.0 

81.9 

655.2 

99  92 

Skeleton  of  chuck  and  neck 

13,202 

5.864 

4222.4 

2012.6 

469.3 

3385.5 

555.01 

Skeleton  of  round 

1530.1 

1756.9 

184.3 

1360.7 

244.00 

Skeleton  of  loin 

6,506 

5,050 

1732.6 

1749.5 

186.5 

1698.2 

242 . 28 

Skeleton  of  rib 

1448.0 

910.1 

169.8 

1451.3 

216.24 

Horns 

1,800 

1,490 

712 

665 . 7 

10.0 

92.2 

350.0 

63.36 

Hoof 8 and  dewclaws 

811.1 

17.0 

104.3 

30.5 

2.43 

Teeth 

188.9 

7.3 

14.0 

403.8 

77.53 

60 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  38. — Steer  509.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

18.291 

14276.7 

627.2 

125.5 

4.21 

4.37 
5.78 
0.54 
2.80 

23.39 

2.38 
1.60 

12.63 

2,616 

3,283 

1829.8 

320.8 

67  8 

24  1 

2529.0 

94.5 

98.6 

33.2 

1,248 

247.2 

949.7 

9.0 

3 7 

739 

494.2 

130.5 

12.4 

11  7 

Digestive  and  excretory  system  (partial) .... 

19.016 

9,922 

13883.8 

1127.1 

2260.2 

8521.8 

419.5 

53.0 

172.7 

17.4 

630 

316.9 

234.0 

11  4 

6.6 

3.875 

110 

2650.8 

68.1 

119.7 

51.7 

Gail 

100.7 

0.1 

0.3 

1.0 

0 03 

1,304 

562 

1007.4 

27.7 

39  0 

18.5 

3.16 
1.51 
1.81 
17.30 
4.82 
19.68 
46  10 

308.6 

159.9 

12.6 

6.6 

774 

596.1 

29.6 

20.5 

8.6 

Hair  and  hide 

37,614 

3,212 

22180.6 

931.7 

2358.4 

385.2 

Head  and  tail , lean  and  fat 

2144.6 

418.3 

98.8 

26  4 

Shin  and  shank,  lean  and  fat 

11.782 

8001.4 

1150.5 

398.2 

107  3 

Flank  and  plate,  lean  and  fat 

31.790 

17051.8 

9195.6 

832.9 

233  0 

Rump,  lean  and  fat 

7.370 

4113.4 

1960.4 

193.7 

59  3 

10.76 

99.89 

79.14 

Chuck  and  neck,  lean  and  fat 

60.176 

41216.4 

5931.0 

1851.0 

538  6 

Round,  lean 

40,376 

29735.7 

1688.9 

1301.3 

405  8 

Round,  fat 

5,106 

30,836 

1305.0 

3269.9 

82.8 

17.5 

2.09 

Loin,  lean 

21471.7 

2778.3 

956.8 

293.3 

55.50 

Loin,  fat 

7,570 

1230.8 

5871.4 

78.2 

19.2 

3 10 

Rib,  lean 

16  360 

11091.3 

1774.1 

478.9 

149.7 

27.81 

0.89 

Rib,  fat 

1,978 

347.2 

1503.7 

20.9 

5.5 

Kidney,  fat 

1.576 

86.1 

1456.4 

5.0 

2.2 

0 27 

Skeleton  of  feet 

6,144 

2523.7 

662.2 

225.4 

1441.2 

260.81 

Skeleton  of  head 

8,247 

3920.6 

699.2 

261.9 

1745.4 

311.08 

Skeleton  of  tail 

386 

146.5 

102.5 

11.1 

58.3 

10  55 

Skeleton  of  shin 

5,046 

1430 . 6 

918.5 

193.7 

1480.5 

260  78 

Skeleton  of  shank 

5.498 

1569.7 

1240.2 

193.5 

1437.4 

276  38 

Skeleton  of  flank  and  plate 

5,124 

2119.7 

667.8 

181.0 

1146.9 

209  16 

Skeleton  of  rump 

2,860 

789.2 

850.7 

87.5 

636.2 

110.91 

Skeleton  of  chuck  and  neck 

13,682 

5,442 

4431.2 

2366.0 

506.8 

3426.8 

630 . 19 

Skeleton  of  round  (excl.  marrow) 

1521.4 

1342.0 

161.5 

1412.1 

251.64 

Marrow  from  skeleton  of  round 

578 

67.4 

502.0 

1.2 

2.1 

0.34 

Skeleton  of  loin 

6,872 

1891.0 

1763.4 

215.1 

1762.5 

303.33 

Skeleton  of  rib 

4,986 

1,590 

1398.6 

837.2 

180.6 

1524.2 

271.44 

Hoofs  and  dewciaws 

1065.0 

7.3 

82.6 

23.2 

1.86 

Teeth 

838 

239.1 

4.5 

14.6 

469.3 

88.27 

Table  39. — Steer  512.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood  

24,176 

19328.5 

13.3 

742.9 

191.0 

5.56 

Circulatory  system 

2'432 

985.7 

1167.0 

41.6 

13.7 

1.34 

Lean  heart 

1,555 

1202.3 

54.0 

43.6 

19.7 

3.34 

Respiratory  system 

3,881 

2929.7 

181.2 

108.8 

41.9 

6.36 

Fat  from  thoracic  cavity 

1.171 

143.2 

1001.5 

3.1 

1.7 

0.18 

Brain  and  spinal  cord 

666 

479.9 

74.0 

11.2 

12.5 

2.56 

Digestive  and  excretory  system  (partial) .... 
Offal  fat 

20.735 

17,454 

15278.4 

1956.9 

2133.2 
15058 . 1 

452.7 

64.4 

157.8 

25.1 

24.67 

3.32 

Heart  and  neck  sweetbreads 

511 

206.4 

252.5 

7.7 

3.8 

0.86 

Liver 

4,416 

212 

3046.3 

115.9 

144.1 

70.1 

14.75 

Gall 

197.5 

0.1 

0.5 

2.2 

0.06 

Spleen 

1,255 

968.2 

29.7 

35.1 

16.8 

3.00 

Pancreas 

736 

422.4 

196.1 

15.4 

9.8 

2.02 

Kidneys 

1,074 

1.766 

829.5 

73.4 

22.3 

11.3 

2.18 

Tongue,  marketable 

1172.3 

237.1 

45.6 

15.9 

2.84 

Hair  and  hide 

41.268 

23189.7 

1490.6 

2701.8 

480.0 

19.40 

Head  and  tail,  lean  and  fat 

4.412 

2715.8 

841.5 

127.6 

37.9 

6.13 

Shin  and  shank,  lean  and  fat 

12.706 

8717.1 

1191.8 

424.8 

117.8 

20.46 

Flank  and  plate,  lean  and  fat 

48,946 

20515.2 

22190.7 

934.9 

279.0 

46.01 

Rump,  lean  and  fat 

10.484 

73,512 

4675.7 

4385.4 

212.5 

69.9 

12.48 

Chuck  and  neck,  lean  and  fat 

46450.8 

13372.6 

2077.5 

675.6 

110.27 

Round,  lean 

43.408 

31805.9 

1978.1 

1405.1 

444.5 

83.34 

Round,  fat 

9,940 

2189.8 

7023.4 

76.0 

30.9 

3.98 

Loin,  lean 

32,062 

15,308 

16,908 

5,398 

21676.2 

3539.6 

986.2 

292.4 

54.51 

Loin,  fat 

1913.0 

12759.8 

99.5 

27.6 

3.98 

Rib,  lean 

11010.3 

2527.8 

501.7 

151.5 

26.55 

Rib,  fat 

806.4 

4338.2 

45.3 

11.4 

1.89 

Kidney,  fat 

4,740 

212.5 

4451.6 

8.7 

6.2 

0.95 

Skeleton  of  feet 

7,016 

9,665 

416 

2627.6 

1004.3 

252.4 

1709.5 

309.05 

Skeleton  of  head 

4169.7 

1252.1 

312.1 

2166.9 

406.32 

Skeleton  of  tail 

153.8 

101.0 

13.0 

75.3 

13.58 

Studies  In  Animal  Nutrition — III 


61 


Table  39. — Steer  512.  Weights  of  Constituents  in  Samples,  Grams — Cont. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Skeleton  of  shin 

6,074 

1649.6 

1263.9 

220.6 

1821.2 

327.27 

Skeleton  of  shank 

6.156 

1968.6 

1326.1 

217.9 

1401.6 

252.03 

Skeleton  of  flank  and  plate 

7,788 

2865.4 

1640.2 

236.2 

1523.6 

279.28 

Skeleton  of  rump 

3.264 

772.9 

1001.4 

97.5 

857.6 

145.12 

Skeleton  of  cnuck  and  neck 

16.536 

4758.2 

3139.5 

536.4 

5045.1 

899.23 

Skeleton  of  round  (excl.  marrow) 

7,430 

2134.1 

1986.3 

209.7 

1679.6 

301 . 14 

Marrow  from  skeleton  of  round 

396 

39.9 

349.7 

0.7 

2.6 

0.52 

Skeleton  of  loin 

8,748 

2198.9 

2134.8 

261.3 

2330.6 

466.62 

Skeleton  of  rib 

6,938 

2042.3 

1132.6 

241.1 

1991.1 

378.40 

Horns 

1.810 

637.6 

8.7 

127.3 

441.7 

70.12 

Hoofs  and  dewclaws 

1.724 

843.1 

10.1 

135.5 

48.1 

2.14 

Teeth 

710 

141.4 

5.6 

15.3 

452.4 

85.42 

Table  40. — Steer  513.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

25.680 

19947.2 

881.1 

185.4 

7.19 

3,485 

4,858 

3.942 

2704.4 

156.8 

93.2 

31.7 

5.47 

3590.3 

341.6 

133  2 

48.2 

7.53 

529.8 

3299 . 7 

14  4 

7.3 

0.95 

748 

510.1 

115  9 

12  9 

12.0 

3.07 

Digestive  and  excretory  system  (partial) .... 
Offal  fat 

25,642 

53,771 

19310.7 
3055  8 

1973.7 

49939.3 

592.3 

115.6 

266.9 
42  5 

40.26 

5.91 

Heart  and  neck  sweetbreads 

1.334 

538.0 

674.3 

18.7 

11.0 

2.64 

Liver 

5,920 

37 

4021.2 

185.1 

190.9 

82.7 

19.71 

Gall 

33.8 

0.1 

0.1 

0.4 

0.01 

Spleen 

1,114 

842.7 

50.6 

32.7 

14.0 

2.70 

Pancreas 

873 

437.5 

302.6 

16.5 

9.5 

2.11 

Kidneys 

1,015 

45,286 

5,390 

17,798 

115.774 

772.1 

58.9 

25.4 

11.5 

2.23 

Hair  and  hide 

26069 . 3 

4561.2 

2344.9 

426.1 

24.45 

Head  and  tail,  lean  and  fat 

3038.8 

1417.1 

131.5 

41.5 

6.90 

Shin  and  shank,  lean  and  fat 

9924.0 

4922.4 

411.0 

124.1 

21.00 

Flank  and  plate,  lean  and  fat 

33830.3 

72079.7 

1437.9 

445.7 

75.25 

Rump,  lean  and  fat 

19,082 

6025.1 

11211.6 

263.0 

83.6 

15.27 

Chuck  and  neck,  lean  and  fat 

110,940 

53117.0 

41259.7 

2387.4 

697.8 

128.69 

Round,  lean 

50.782 

19.108 

33089.0 

7280.6 

1473.2 

462.8 

87.85 

Round,  fat 

3394.2 

14539 . 1 

176.0 

38.0 

4.59 

Loin,  lean 

44,510 

49.928 

26437.2 

9612.8 

1236.0 

367.7 

72.55 

Loin,  fat 

4444.0 

44182.3 

182.7 

57.4 

8.99 

Rib,  lean 

24,744 

13754.9 

6857.3 

635.2 

187.3 

35.14 

Rib,  fat 

23.608 

4201  5 

17813.4 

96  3 

32.6 

4.96 

Kidney,  fat 

14.490 

566.9 

13755.1 

22  6 

10.7 

1.45 

Skeleton  of  feet 

7.598 

2753.4 

1128  8 

267  6 

1811.1 

322.38 

Skeleton  of  head 

7,865 

3201.6 

742.5 

259.6 

1968.5 

460.57 

Skeleton  of  tail 

297 

116.7 

58.7 

10.0 

51.2 

9.00 

Skeleton  of  shin 

6.120 

1792.6 

1091.5 

206.7 

1672.3 

294.43 

Skeleton  of  shank 

6.058 

1582.6 

1206.7 

216.3 

1792.6 

317.02 

Skeleton  of  flank  and  plate 

7.438 

3,464 

15.526 

3043 . 8 

1121.7 

243.8 

1519.2 

307.86 

Skeleton  of  rump 

845.8 

1014.5 

104.2 

855.2 

162.22 

Skeleton  of  chuck  and  neck 

4969.6 

2760.7 

544.7 

3912.9 

785.31 

Skeleton  of  round  (excl.  marrow) 

6,564 

480 

1625.0 

2011.1 

194.8 

1625.1 

286.58 

Marrow  from  skeleton  of  round 

41.6 

431.8 

0.8 

3.9 

0.67 

Skeleton  of  loin 

8,536 

2149.9 

2543.6 

258.2 

2081.3 

401.11 

Skeleton  of  rib 

7.096 

1884.6 

1528.4 

238.6 

2073.2 

392.27 

Horns 

2.144 

790  1 

16  4 

138  3 

485.0 

89  36 

Hoofs  and  dewclaws 

2.180 

914.1 

19.4 

195.2 

64.5 

5.21 

Teeth 

874 

278.8 

9 6 

15.8 

459.5 

87.04 

62 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  41. — Steer  515.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

27,856 

22036.1 

896.1 

165.5 

6.69 

Circulatory  system 

5,787 

2432.4 

2808.4 

73.8 

27.8 

4.98 

Respiratory  system 

3,653 

2789.2 

198.8 

98.6 

35.4 

6.83 

Brain  and  spinal  cord 

728 

509.9 

117.1 

12.0 

11.9 

2.88 

Digestive  and  excretory  system 

36,311 

24025.9 

6794.2 

733.1 

309.0 

58.82 

Offal  fat 

29,877 

2250.3 

26976.2 

84.0 

37.1 

4.48 

Hair  and  nide 

49.943 

27969.6 

6267.9 

2417.2 

927.4 

29.47 

Head  and  tail,  lean  and  fat 

5.918 

3169.8 

1807.2 

143.0 

43.3 

7.70 

Shin  and  shank,  lean  and  fat 

16,676 

9565.5 

4461.3 

417.2 

119.4 

20.51 

Flank  and  plate,  lean  and  fat 

87.138 

27115.6 

52220.7 

1185.1 

338.1 

61.87 

Rump,  lean  and  fat 

15,810 

5586.6 

8569.7 

238.1 

75.3 

13.44 

Chuck  and  neck,  lean  and  fat 

88,134 

46944.6 

27399.1 

2076.4 

627.5 

111.05 

Round,  lean 

42,942 

29067.9 

4746.0 

1332.1 

396.8 

76.01 

Round,  fat 

19.058 

3339.3 

14807.3 

149.4 

41.0 

4.96 

Loin,  lean 

41,620 

27036.8 

6323.3 

1248.2 

413.7 

75.33 

Loin,  fat 

38,324 

4053.2 

33197.4 

157.9 

44.1 

6.52 

Rib,  lean 

19.016 

11634.6 

3976.8 

527.9 

161.5 

28.90 

Rib,  fat 

16,282 

1476.9 

14374.9 

64.2 

21.8 

3.26 

Kidnev,  fat 

9.922 

491.2 

9369.6 

17.7 

'8.0 

1.49 

Skeleton  of  feet,  head  and  tail 

18.179 

7520.3 

2029.0 

551.2 

4270.3 

739.16 

Skeleton  of  shin  and  shank 

13,900 

3852.1 

3761.8 

423.3 

3429.1 

560.45 

Skeleton  of  flank  and  plate 

6,368 

2687.0 

1016.6 

217.2 

1132.2 

198.24 

Skeleton  of  rump 

4,074 

1213.8 

104.7 

126.1 

949.8 

165.85 

Skeleton  of  chuck  and  neck 

14,528 

4431.0 

2254.3 

484.2 

4295.6 

617.88 

Skeleton  of  round 

6.344 

1499.9 

1866.0 

191.8 

1825.0 

310.22 

Skeleton  of  loin 

7,784 

1969.9 

2196.6 

231.3 

2040.7 

350.05 

Skeleton  of  rib 

6,464 

1686.2 

1303.1 

211.4 

2015.2 

282.74 

Horns 

1,804 

733.2 

10.7 

101.3 

427.1 

76.18 

Hoofs  and  dewclaws 

1,893 

1017.5 

10.0 

139.4 

34.5 

1.36 

Teeto 

786 

214.8 

6.8 

14.0 

499.1 

86.40 

Table  42. — Steer  523.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

15,287 

3,044 

12309.1 

426.5 

100.7 

3.36 

Circulatory  system 

1615.0 

1090.5 

51.3 

18.2 

3.35 

Respiratory  system 

3,371 

2652.3 

136.7 

87.8 

33.2 

6.37 

Brain  and  spinal  cord 

797 

546.7 

140.4 

12.9 

12.1 

2.86 

Digestive  and  excretory  system 

24,667 

18548.1 

2548.1 

540.2 

194.4 

38.97 

Hair  and  hide 

33,097 

20547.0 

380.6 

1854.4 

340.9 

16.88 

Head  and  tail,  lean  and  fat 

3,100 

2113.2 

358.8 

95.9 

28.6 

4.77 

Shin  and  shank,  lean  and  fat 

8,684 

6233.9 

524.2 

295.4 

79.5 

14.68 

Flank  and  plate,  lean  and  fat 

26,984 

16058.2 

6184.7 

726.7 

199.4 

37.51 

Rump,  lean  and  fat 

5,418 

2943.6 

1585.8 

137.2 

40.5 

8.34 

Cnuck  and  neck,  lean  and  fat 

50.320 

35668.8 

5176.9 

1502.6 

463.5 

86.05 

Round,  lean 

33,900 

26078.9 

783.1 

1069.6 

353.2 

68.48 

Loin,  lean 

25,834 

18020.3 

2688.3 

782.0 

237.9 

44.43 

Rib,  lean 

12.032 

8456.3 

1118.6 

375.3 

111.5 

21.42 

Round,  fat 

4,556 

1346.0 

2743.8 

72.0 

16.3 

2.19 

Loin,  fat 

6,376 

1051.9 

4969.1 

55.0 

17.9 

2.87 

Rib,  fat 

1,522 

358.3 

989.0 

23.0 

6.0 

' 0.90 

Kidney,  fat 

3,110 

163.6 

2883.8 

14.5 

5.6 

0.47 

Offal  fat 

7,915 

1220.3 

6433.6 

39.2 

16.2 

2.37 

Skeleton  of  feet,  head  and  tail 

13.120 

5805.7 

1065.3 

467.5 

2990.8 

542.51 

Skeleton  of  shin  and  shank 

8,862 

2611.1 

1804.7 

299.6 

2449.0 

454.80 

Skeleton  of  flank  and  plate 

3,882 

1741.8 

415.8 

134.7 

754.8 

134.59 

Skeleton  of  rump 

1,770 

478.5 

358.6 

57.9 

523.8 

196.11 

Skeleton  of  chuck,  and  neck 

9,786 

3314.1 

1261.8 

360.2 

2876.5 

527.56 

Skeleton  of  round 

4,630 

1845.1 

1231.5 

89.9 

1950.2 

177.79 

Skeleton  of  loin 

5,322 

1483.4 

1260.6 

167.4 

1438.5 

265.09 

Skeleton  of  rib 

3,844 

1200.0 

481.2 

138.1 

1219.1 

221.76 

Horns 

1,167 

539.5 

7.3 

65.5 

218.7 

39.87 

Hoofs  and  dewclaws* 

1.063 

575.1 

7.8 

76.5 

21.6 

3 131 

Teeth 

766 

201.6 

5.9 

16.1 

432.0 

81.78 

♦This  sample  was  lost  before  analysis.  The  analysis  of  the  hoofs  and  dewclaws  of  four  animals  of  the  same 
Group  was  used. 


Studies  In  Animal  Nutrition — III 


63 


Table  43. — Steer  524.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

17,019 

2.953 

13956.6 

474.3 

127.0 

3.74 

1833.5 

662.0 

66.2 

22.7 

3.90 

3455 

758 

2682.3 

89.9 

96.5 

37.1 

6.15 

555.0 

77.7 

12.1 

10.8 

2.56 

Digestive  and  excretory  system  (partial) 

17,924 

5.007 

13280.6 

1255.2 

1760.3 

3477.1 

412.8 

38.1 

158.6 

16.0 

26.89 

1.75 

Heart  and  neck  sweetbreads 

541 

406.4 

40.3 

14.1 

10.1 

2.50 

3,019 

2108  5 

100.1 

92.0 

42.5 

9.87 

Gall 

229 

217  2 

0.2 

0.4 

1.8 

0.07 

Spleen 

596.2 

13.8 

20.8 

10.1 

1.97 

435 

285.5 

69.1 

11  1 

5.4 

1.24 

Kidneys 

766 

588.5 

43.9 

18.6 

8.9 

1.59 

30,092 

17832.2 

545.6 

1895.8 

474.6 

17.45 

Head  and  tail,  lean  and  fat 

3.364 

2229.0 

441.6 

105.2 

34.5 

6.02 

Shin  and  shank,  lean  and  fat 

8.674 

6335.2 

485.8 

253.7 

84.5 

14.40 

Flank  amd  plate,  lean  and  fat 

19,788 

4,030 

46,386 

37,714 

2,526 

24.200 

12607.1 

3041.8 

633.2 

188.4 

30.47 

Rump,  lean  and  fat 

2563.5 

691.0 

119.2 

39.2 

7.05 

Cnuck  and  neck,  lean  and  fat 

33655.4 

2918.6 

1465 . 8 

452.3 

80.71 

Round,  lean 

28988.5 

1024.3 

1228.3 

393.4 

72.41 

Round,  fat 

915.9 

1262.0 

56 . 5 

10.6 

1.34 

Loin,  lean 

17588.6 

1099.7 

801.0 

255.8 

46.95 

Loin,  fat 

2,444 

13,144 

459.7 

1791.9 

30.2 

8.6 

1.30 

Rib,  lean  and  fat 

9238.3 

1044.8 

435.1 

129.1 

23.92 

Kidney,  fat 

766 

87.6 

644.9 

3.4 

1.4 

0.24 

Skeleton  of  feet 

6,010 

2434.5 

781.4 

215.9 

1473.4 

262.94 

Skeleton  of  head  and  tail 

8,318 

3820.5 

864.2 

254.0 

1985.3 

354.60 

Skeleton  of  snin  and  shank 

10,262 

3240 . 8 

1982.3 

345.3 

2659.9 

474.00 

Skeleton  of  flank  and  plate 

5,926 

2569.4 

905.0 

190.3 

1135.7 

198.22 

Skeleton  of  rump 

2,424 

12,896 

5,878 

6,586 

831  6 

520.0 

72.7 

542.9 

98.34 

Skeleton  of  chuck  and  neck 

5197.0 

2043.2 

427.6 

2809.5 

514.03 

Skeleton  of  round 

1825.5 

1665.9 

156.7 

1334.3 

238.82 

Skeleton  of  loin 

1926.5 

1771.7 

187.4 

1600.8 

279.64 

Skeleton  of  rib 

5,310 

1.227 

1897.5 

984.2 

157.7 

1324.9 

235.18 

Horns* 

506.9 

9.1 

78.7 

237.5 

44.07 

Hoofs  and  dewclaws 

1,494 

806 

750.6 

12.4 

112.8 

47.7 

3.27 

Teeth 

217.4 

8.9 

15.4 

466.4 

88.56 

♦This  sample  was  lost  before  analysis.  The  analysis  of  (he  horns  of  a steer  of  the  same  age  was  used. 


Table  44. — Steer  525.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

13,614 

10961.2 

400.5 

89.9 

3.58 

Circulatory  system 

1,320 

846.0 

324.0 

22.5 

9.4 

1.59 

Respiratory  system 

1,658 

1303.4 

43.7 

46.8 

18.4 

3.20 

Brain  and  spinal  cord 

711 

505.2 

99.1 

12.4 

10  7 

2.63 

Digestive  and  excretory  system 

23,001 

17820.9 

2103.2 

459.8 

203.1 

35.95 

Hair  and  hide 

27,813 

17354.2 

138.5 

1592.0 

370.2 

15.85 

Head  and  tail,  lean  and  fat 

2,788 

2072.0 

364.7 

51.8 

27.5 

4 40 

Shin  and  shank,  lean  and  fat 

7,596 

5533.6 

404.9 

253.5 

73.3 

13.44 

Flank  and  plate,  lean  and  fat 

18,762 

11412.4 

3790.7 

539.0 

155.4 

27.96 

Rump,  lean  and  fat 

4.154 

2532.3 

869.2 

114.7 

35.9 

6.73 

Chuck  and  neck,  lean  and  fat 

35,824 

25489.1 

3024.6 

1119.5 

340.0 

63.77 

Round,  lean 

27,524 

21202.8 

661.4 

858.8 

292.9 

56.42 

Loin,  lean 

18.710 

13964.0 

649.1 

605.6 

196.1 

37.42 

Rib,  lean 

11,666 

8226.2 

1010.2 

368.8 

114.0 

21.12 

Round,  fat 

1,962 

641.3 

1114.9 

31.4 

8.5 

1.10 

Loin,  tat 

3.758 

826.8 

2615.2 

41.3 

11.1 

1.50 

Rib,  fat 

664 

186.5 

409.1 

10.7 

4.0 

0.61 

Kidney,  fat 

1,258 

84.6 

1134.5 

5.9 

2.0 

0.25 

Offal  tat 

4,961 

1043.2 

3504.5 

63.8 

14.5 

2.53 

Skeleton  of  feet,  head  and  tail 

10,782 

4638.5 

1077  1 

402.8 

2352 . 9 

4*0.32 

Skeleton  of  shin  and  shank 

7,014 

2127.6 

1342.1 

244.4 

2071.0 

293.89 

Skeleton  of  flank  and  plate 

3,454 

1540.9 

374  9 

116.2 

744.9 

103.24 

Skeleton  of  rump 

1,542 

474.9 

367.3 

49.9 

379.5 

65.01 

Skeleton  of  chuck  and  neck 

8,450 

2885.3 

1665.8 

261.3 

1935.1 

363.43 

Skeleton  of  round 

4,046 

1157.2 

1322.5 

107.3 

881.3 

114.34 

Skeleton  of  loin 

3,928 

1123.2 

994.1 

118.6 

959  8 

144.47 

Skeleton  of  rib 

3,888 

1233.9 

701.8 

139.7 

961  9 

169.21 

Horns 

1,298 

634.4 

7.1 

73.2 

217.0 

40  77 

Hoofs  and  dewciaws 

940 

487.9 

5.4 

74.2 

11.8 

1.26 

Teeth 

690 

159.6 

7.3 

13.1 

418  9 

80.99 

64 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  45. — Steer  526.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

18,957 

15151.6 

585.2 

147.3 

4 74 

Circulatory  system 

2,413 

1527.8 

517.2 

56.5 

19.6 

3.50 

Respiratory  system 

3,797 

2909.1 

124.9 

106.3 

38.9 

5.92 

Fat  from  the  thoracic  cavity 

1.585 

340.3 

1159.3 

11.9 

4.5 

0.54 

Brain  and  spinal  cord 

660 

483.2 

67.7 

11.5 

11.4 

2.77 

Digestive  and  excretory  system  (partial) 

20,541 

14971.1 

2510.5 

437.5 

157.8 

26.09 

Offal  fat 

11,551 

1500.7 

9719.5 

47.4 

20.2 

2.08 

Heart  and  neck  sweetbreads 

439 

265.5 

106.3 

9.9 

6.3 

1.53 

Liver 

3,531 

2398.3 

150.2 

114.7 

51.9 

12.29 

Gall 

143 

132.1 

0.2 

0.3 

1.7 

0.04 

Spleen 

831 

652.3 

12.4 

23.0 

12.9 

2.47 

Pancreas 

498 

316.0 

90.7 

11.2 

5.8 

1.40 

Kidneys 

922 

678.2 

83.6 

21.7 

9.9 

1.83 

Hair  and  hide 

35,732 

20663.1 

2041.7 

2116.4 

514.5 

17.87 

Head  and  tail,  lean  and  fat 

3.616 

2234.4 

710.2 

105.2 

30.1 

5.13 

Shin  and  shank,  lean  and  fat 

11,644 

8261.4 

877.6 

384.4 

104.8 

19.10 

Flank  and  plate,  lean  and  fat 

39.524 

19561.2 

13931.8 

920.9 

268.0 

49.41 

Rump,  lean  and  fat 

8,594 

4611.8 

2576.6 

211.7 

63.3 

11.9 

Chuck  and  neck,  lean  and  fat 

61.228 

40068.2 

8878.1 

1789.7 

555.3 

101.64 

Round,  lean 

44,614 

29839.6 

5303.3 

1472.3 

460.0 

85.21 

Round,  fat 

5.016 

1136.9 

3460.1 

83.8 

14.3 

1.81 

Loin,  lean 

31,440 

22620.5 

1679.2 

1009.2 

320.1 

59.74 

Loin,  fat 

11,634 

1687.9 

9298.9 

98.9 

24.2 

3.49 

Rib,  lean 

17,264 

12022.8 

1728.8 

531.2 

158.8 

28.49 

Rib.  fat 

3,720 

641.4 

2781.4 

43.2 

9.6 

1.64 

Kidney,  fat 

3,224 

292.6 

2831.7 

11.0 

4.4 

0.68 

Skeleton  of  feet 

6.138 

2358.5 

850.3 

226.0 

1440.9 

267.25 

Skeleton  of  head  and  tail 

9.165 

4249.2 

1173.6 

274.8 

1925.0 

351.20 

Skeleton  of  shin  and  shank 

11.612 

3228.4 

2333.7 

377.9 

3333.0 

608.24 

Skeleton  of  flank  and  plate 

6,588 

2645.4 

1084.8 

209.2 

1339.5 

249.88 

Skeleton  of  rump 

2.838 

832.8 

726.8 

88.2 

672.8 

121.38 

Skeleton  of  chuck  and  neck 

13,842 

4749.6 

2415.3 

462.2 

3405.3 

620.40 

Skeleton  of  round 

6,352 

17! S. 7 

1922.9 

171.1 

1478.2 

274.28 

Skeleton  of  loin 

6 344 

1995.9 

1939.1 

193.4 

1595.6 

293.06 

Skeleton  of  rib 

5.690 

1804.4 

1142.3 

188.6 

1495.7 

270.73 

Horns 

1.427 

589.6 

10.6 

91.5 

276.2 

51.26 

Hoofs  and  dewclaws 

1.875 

1019.9 

11.7 

136.8 

39.9 

1.59 

Teeth 

782 

173.2 

10.0 

15.2 

480.7 

91.39 

Table  46. — Steer  527.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood  . 

27,382 

4,903 

21615  6 

899  0 

208  7 

6 30 

Circulatory  system 

2645.2 

1608.1 

94.5 

31.1 

5.59 

Respiratory  system 

4.326 

3172.3 

310.9 

117.5 

44.3 

7.18 

Fat  from  thoracic  cavity 

Brain  and  spinal  cord 

3.988 

701 

412.4 

490.4 

3495.3 

98.6 

13.4 

11.1 

5.1 

10.8 

0.60 

2.58 

Digestive  and  excretory  system  (partial) 

Offal  fat 

23,818 

48,517 

1.037 

15915.6 

2617.0 

4412.8 

45247.0 

494.7 

88.8 

169.1 

57.3 

28.34 

6.31 

Heart  and  neck  sweetbreads 

379.8 

574.0 

12.3 

7.6 

3.19 

Liver 

5.720 

3882.3 

198.6 

188.4 

90.1 

10.52 

Spleen 

1,226 

941.5 

26.6 

35.6 

15.7 

2.91 

Pancreas 

849 

353.0 

393.1 

12.4 

7.5 

1.72 

Kidneys 

1,244 

937.3 

103.0 

27.5 

12.5 

2.40 

Hair  and  hide 

46.240 

25189.2 

5483.6 

2458.6 

633.0 

25.89 

Head  and  tail,  lean  and  fat 

5,018 

2705.8 

1537.3 

120.9 

34.7 

6.17 

Shin  and  shank,  lean  and  fat 

17,358 

118,978 

24.020 

9807.8 

4582.2 

446.1 

130.9 

22.57 

Flank  and  plate,  lean  and  fat 

32362.0 

77919.9 

1305.2 

371.2 

70.20 

Rump,  lean  and  fat 

6792.1 

15123.5 

296.2 

75.7 

14.17 

Chuck  and  neck,  lean  and  fat 

112,440 

52601.7 

44206.9 

2282.5 

693.8 

122.56 

Round,  lean 

51.396 

33977.4 

7160.0 

1538.3 

444.1 

89.94 

Round,  fat 

21.466 

3462.3 

17027.5 

161.6 

43.8 

4.72 

Loin,  lean 

50,140 

30681.7 

10276.7 

1402.4 

425.7 

80.73 

Loin,  fat 

52,724 

5018.8 

46540.0 

188.2 

67.0 

8.96 

Rib,  lean 

25.860 

14218.9 

7266.1 

658.7 

203.5 

36.46 

Rib,  fat 

24.278 

2297.4 

21415.1 

98.8 

32.3 

3.88 

Kidney,  fat 

18,964 

1028.4 

17679.6 

35.5 

19.3 

2.65 

Skeleton  of  feet 

7.442 

2774.4 

1199.3 

250.9 

1761.1 

286.44 

Skeleton  of  head  and  tail 

8'S22 

3717.2 

2189.8 

279.2 

1913.7 

324.30 

Skeleton  of  shin  and  shank 

13.136 

3580.4 

2794.8 

413.7 

3726.6 

597.69 

Skeleton  of  flank  and  plate 

6 082 

2392.3 

1121.3 

182.2 

1187.9 

207.03 

Skeleton  of  rump 

3.260 

810.0 

1000.5 

96.7 

780.0 

122.15 

Skeleton  of  chuck  and  neck 

14,870 

4172.2 

3653.6 

458.3 

3723.0 

586.47 

Skeleton  of  round 

6.446 

1253.5 

2188.7 

170.3 

1742.5 

319.98 

Skeleton  of  loin 

7.140 

1850.7 

1840.8 

225.0 

1924.2 

368.14 

Skeleton  of  rib 

6,546 

1762.2 

1510.4 

197.8 

1873.6 

360.23 

Horns 

1.266 

451.5 

9.8 

88.0 

250.8 

46.31 

Hoofs  and  dewclaws 

2.174 

958.3 

20.9 

193.8 

44.8 

3.15 

Teeth 

872 

180.5 

12.5 

17.0 

552.7 

120.61 

Studies  In  Animal  Nutrition — III 


65 


Table  47. — Steer  531.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

9,457 

7743.6 

268.7 

66.5 

3.78 

Circulatory  system 

1,971 

1118.0 

577.9 

38.8 

13.3 

2.48 

Respiratory  system 

1,915 

1500.2 

54.0 

51.3 

20.2 

3.91 

Brain  and  spinal  cord 

573 

418.2 

77.3 

10.1 

8.6 

2.08 

Digestive  and  excretory  system  (partial) 

11,807 

9106.6 

1089.8 

233.4 

79.0 

14.29 

Offal  fat 

2.899 

751.7 

2049.0 

18.0 

8.4 

1.33 

Heart  and  neck  sweetbreads 

472 

320.9 

69.8 

11.8 

8.1 

2.02 

Liver 

2,205 

1582.9 

42.9 

64.4 

30.5 

7.08 

Gall 

86 

78.8 

0.2 

0.8 

0.04 

Spleen 

481 

361.1 

20.2 

14.2 

6.9 

1.40 

Pancreas 

297 

199.2 

43.9 

7.5 

3.7 

0.86 

Kidneys 

506 

379.9 

41.9 

11.5 

5.3 

1.08 

Hair  and  Hide 

16,693 

10627.6 

135.4 

957.0 

189.6 

12.52 

Head  and  tail,  lean  and  fat 

1,722 

1157.5 

242.3 

50.3 

15.0 

2.67 

Shin  and  shank,  lean  and  fat 

5,480 

3934.2 

259.5 

197.1 

58.7 

10.30 

Flank  and  plate,  lean  and  fat 

10.854 

6553.0 

1865.9 

331.6 

109.1 

19.10 

Rump,  lean  and  fat 

2,506 

1553.5 

453.9 

74.7 

24.5 

4.43 

Chuck  and  neck,  lean  and  fat 

26,902 

18808.4 

2099.7 

880.8 

278.2 

52.73 

Round,  lean 

21.496 

16145.9 

393.6 

703.4 

236.9 

44.71 

Round,  fat 

1,490 

424.3 

904.2 

22.7 

7.1 

1.06 

Loin,  lean 

14,078 

10278.5 

602.3 

466.3 

154.9 

28.72 

Loin,  fat 

2.264 

560.5 

1468.5 

31.4 

9.9 

1.70 

Rib,  lean  and  fat 

6,612 

4597.7 

500.0 

220.7 

69.6 

12.56 

Kidney  fat 

726 

44.0 

655.3 

4.3 

1.7 

0.23 

Skeleton  of  feet 

3,762 

1496.3 

540.0 

121.6 

927.0 

166.73 

Skeleton  of  head  and  tail 

4.842 

2376.4 

390.8 

147.3 

1009.5 

175.72 

Skeleton  of  shin  and  shank 

5,998 

1903.2 

1100.4 

176.2 

1644.4 

306.68 

Skeleton  of  flank  and  plate 

2,546 

1190.8 

269.1 

82.0 

519.3 

88.19 

Skeleton  of  rump 

1.060 

333.0 

225.1 

34.7 

270.7 

48.47 

Skeleton  of  chuck  and  neck 

6.098 

2286.3 

976.6 

215.6 

1434.6 

255.32 

Skeleton  of  round 

3,640 

1275.1 

943.2 

97.5 

777.6 

138.54 

Skeleton  of  loin 

2,834 

881.1 

621.1 

93.5 

722.5 

126.71 

Skeleton  of  rib 

2.280 

738.4 

381.7 

85.2 

575.2 

104.58 

Hoofs  and  dewclaws 

790 

411.6 

6.5 

59.9 

15.5 

0.99 

Teeth 

426 

116.8 

3.8 

8.5 

237.7 

45.26 

Table  48. — Steer  532.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

18,752 

15090.3 

562.0 

122.3 

6.19 

Circulatory  system 

4,843 

2320.4 

1949.6 

77.7 

27.4 

5.09 

Respiratory  system 

3,870 

3001.2 

132.1 

102.0 

40.7 

7.47 

Brain  and  spinal  cord 

643 

465.9 

93.0 

10.7 

10.4 

2.52 

Digestive  and  excretory  system  partial 

21,741 

15805.9 

3039.2 

412.6 

153.9 

29.13 

Offal  fat 

23,697 

2263.5 

21044.1 

57.6 

27.5 

3.79 

Heart  and  neck  sweetbreads 

441 

218.1 

169.6 

7.2 

4.4 

1.05 

Liver 

5,694 

4066.6 

135.2 

161.9 

78.3 

17.48 

Gall  

185 

170.7 

0.4 

1.9 

0.08 

Spleen 

884 

660.3 

44.5 

25.5 

11.1 

2.32 

Pancreas 

630 

353.8 

174.4 

12.8 

7.4 

1.85 

Kidneys 

868 

618.4 

95.3 

21.9 

8.8 

1.85 

Hair  and  hide 

33,988 

20244.6 

2324.4 

1775.9 

350.8 

24.13 

Head  and  tail,  lean  and  fat 

4,280 

2571.8 

935.1 

114.5 

34.4 

6.21 

Shin  and  shank,  lean  and  fat 

12.008 

8109.7 

1320.6 

379.5 

112.0 

20.05 

Flank  and  plate,  lean  and  fat 

44,636 

19529.6 

18635.1 

998.5 

292.4 

51.78 

Rump,  lean  and  fat 

8.058 

3910.4 

2891.5 

189.0 

57.7 

10.72 

Chuck  and  neck,  lean  and  fat 

66.204 

40908.8 

12261.0 

1926.5 

595.2 

103.29 

Round,  lean 

38.064 

27369.5 

1988.1 

1257.6 

405.4 

76.13 

Round,  fat 

6,064 

1316.7 

4264.3 

67.5 

17.9 

2.55 

Loin,  lean 

36,136 

24719.2 

3466.5 

1175.1 

362.8 

66.49 

Loin,  fat 

14,954 

1872.2 

12472.4 

100.8 

28.6 

4.93 

Rib,  lean 

17,356 

11677.1 

1993.0 

547.8 

169.9 

30.55 

Rib,  fat 

6.194 

967.6 

4885.7 

52.8 

16.6 

2.48 

Kidney, fat 

11,734 

383.8 

11174.1 

17.6 

10.0 

2.58 

Skeleton  of  feet 

6,490 

2536.3 

031  3 

234 . 1 

1547.5 

273.94 

Skeleton  of  head  and  tail 

7.120 

3304.3 

910.4 

212.5 

1512.7 

270.13 

Skeleton  of  shin  and  shank 

10,756 

3107.4 

2325.1 

464.2 

2512.3 

472.94 

Skeleton  of  flank  and  plate 

5,478 

2132  7 

1037.9 

165.8 

819.4 

149.99 

Skeleton  of  rump 

2.282 

659 . 5 

633.1 

73.1 

517.8 

93.20 

Skeleton  of  chuck  and  neck 

13,014 

3911  0 

3071  4 

438.2 

3126.9 

506.24 

Skeleton  of  round 

5.024 

1511.0 

1914.0 

141  4 

1169.5 

208.82 

Skeleton  of  loin 

6.246 

17.51  9 

1827.4 

193.2 

1377.4 

254.46 

Skeleton  of  rib 

5,222 

1826.1 

1175.4 

154.1 

1163.5 

209.35 

Horns 

228 

124.8 

1.2 

14  4 

17.3 

3.28 

Hoofs  and  dewclaws 

1,406 

730.2 

9.1 

107.0 

27.4 

2.15 

Teeth 

494 

153.2 

4.4 

10.6 

25  V 4 

50.16 

66 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  49. — Steer  538.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

7,219 

1,670 

1,825 

487 

10,290 

3,452 

481 

5948.5 

7.0 

197.4 

56  7 

2.02 

926.6 

479.3 

32.3 

10  6 

1 90 

Respiratory  system 

Brain  and  spinal  cord 

Digestive  and  excretory  system  (partial) . . . 
Offal  fat 

1474.6 
362.0 

7822.7 
731.2 

37.5 

57.2 

1009.8 

2639.2 

45.7 
7.8 

217.2 

18.8 

18.6 

7.3 
89.2 

9.4 
8 0 

3.63 
1.69 
15.54 
1.90 
1.95 
6.23 
0 09 

335.8 

70.2 

11.2 

1,978 

122 

1394.1 

37.9 

58.5 

27  7 

Gall 

110.9 

0.1 

0.3 

1 3 

331 

259.6 

5.1 

9.5 

4.7 

0.90 

0.73 

208 

144.9 

24.2 

5.3 

3.1 

487 

356.6 

54.9 

11.2 

5 2 

1.02 

15,342 

1,496 

9870.9 

205.9 

853.6 

163.1 

9 67 

Head  and  tail,  lean  and  fat. 

991.9 

231  0 

40.4 

13  4 

2.21 
7 50 

Shin  and  shank,  lean  and  fat 

4,190 

11,036 

3008.9 

241.9 

138.0 

42  0 

Flank  and  plate,  lean  and  fat 

6367.8 

2589.7 

306.1 

94.0 

17.66 

Rump,  lean  and  fat 

2,096 

1252.3 

442.6 

55.9 

18.7 

3.23 

Chuck  and  neck,  lean  and  fat 

22.284 

15261.6 

2665.8 

662.1 

207.5 

38  77 

Round,  lean 

16,324 

1,702 

12401.5 

440.6 

515.7 

178.4 

32.97 

Round,  fat 

519.6 

999.6 

29.1 

7.6 

1.16 

Loin,  lean 

12.732 

9312.3 

715.2 

396.9 

134.3 

25  85 

Loin,  fat 

2.360 

5,196 

470.5 

1713.4 

23.1 

6.9 

1.27 

Rib,  lean 

3683.0 

423.5 

159.8 

52.8 

9 92 

Rib, fat 

'426 

128.6 

240.4 

6.2 

3.1 

0 49 

Kidney, fat 

622 

42.0 

565.8 

2.1 

1.1 

0.21 

Skeleton  of  feet 

3,166 

4,001 

135 

1350.6 

491.6 

104.3 

589.4 

105.81 

Skeleton  of  head 

1991.7 

311.8 

116.7 

863.4 

163.60 

Skeleton  of  tail 

70.8 

19.1 

4.3 

15.2 

2.63 

Skeleton  of  shin 

2,208 

2,608 

2.144 

732 

632.3 

526.1 

62.4 

554.4 

86.64 

Skeleton  of  shank 

792.0 

585.2 

88.4 

616.9 

95.71 

Skeleton  of  flank  and  plate 

1054.6 

311.8 

68.6 

322.6 

52.46 

Skeleton  of  rump 

250.9 

173.8 

22.3 

162.7 

27.79 

Skeleton  of  chuck  and  neck 

5,412 

2.556 

2064.1 

917.4 

180.6 

1297.2 

192.40 

Skeleton  of  round 

765.8 

823.7 

61.9 

526.5 

81.20 

Skeleton  of  loin 

2,696 

942.7 

625.4 

79.3 

573.1 

112.23 

Skeleton  of  rib 

2,154 

250 

783.3 

357.8 

68.5 

487.2 

86.83 

Horns 

137.3 

1.3 

14.5 

24.9 

4.74 

Hoofs  and  dewclaws* 

635 

423.1 

3.0 

34.0 

6.5 

0.43 

Teeth  f 

240 

68.83 

2.7 

4.7 

130.0 

24.48 

Table  50. — Steer  540.  Weights  of  Constituents  in  Samples,  Grams. 


Blood 

6,967 

1,436 

1,501 

512 

5732.5 

70.5 

195.5 

50.4 

1.81 

Circulatory  system 

834.8 

398.2 

29.3 

9.7 

1.82 

Respiratory  system 

1194.9 

34.7 

38.9 

15.5 

2.97 

Brain  and  spinal  cord 

377.1 

61.1 

8.3 

8.1 

1.92 

Digestive  and  excretory  system  (partial) 

Offal  fat 

9,280 

2.307 

6982.2 

582.4 

998.2 

1639.3 

185.1 

12.6 

65.2 

6.5 

11.97 

1.08 

Heart  and  neck  sweetbreads 

408 

280.7 

63.0 

9.2 

6.2 

1.52 

Liver 

1,593 

58 

1124.5 

28.0 

45.2 

22.5 

4.86 

Gall 

54.4 

0.04 

0.1 

0.7 

0.02 

Spleen 

331 

257.5 

4.5 

10.3 

4.7 

0.90 

Pancreas 

180 

130.0 

17.3 

4.6 

2.6 

0.62 

Kidneys 

363 

261.9 

38.1 

8.6 

3.9 

0.81 

Hair  and  hide 

12.994 

8399.6 

305.8 

666.2 

163.2 

8.71 

Head  and  tail,  lean  and  fat 

1.274 

866.2 

170.8 

34.9 

11.3 

2.13 

Shin  and  shank,  lean  and  fat 

3,762 

2793.8 

1*1.9 

127.8 

39.1 

6.96 

Flank  and  plate,  lean  and  fat 

8,824 

1,964 

5375.9 

1629.0 

266.6 

77.9 

13.94 

Rump,  lean  and  fat 

1198.9 

384.8 

54.1 

18.1 

3.26 

Chuck  and  neck,  lean  and  fat 

17,978 

13,456 

12783.8 

1500.6 

532.3 

177.3 

32.00 

Round,  lean 

1018". 9 

303.4 

431.7 

145.6 

28.26 

Round,  fat 

810 

225.4 

504.7 

11.0 

3.1 

0.49 

Loin,  lean 

10,700 

2.308 

7869.3 

448.0 

340.2 

115.0 

21.51 

Loin,  fat 

452.3 

1700.8 

22.1 

6.7 

1.34 

Rib,  lean  and  fat 

5,046 

3503.4 

488.3 

156.2 

51.1 

9.14 

Kidney  fat 

Skeleton  of  feet 

682 

2.784 

91.4 

1225.2 

544.3 

337.6 

7.9 

91.5 

1.3 

580.2 

0.24 

99.47 

Skeleton  of  head  

3,682 

138 

1903.2 

259.8 

17.2 

101.4 

788.6 

149.34 

Skeleton  of  tail 

76.4 

3.1 

13.9 

2.44 

Skeleton  of  shin 

1,952 

618.5 

432.5 

68.3 

472.0 

89.03 

Skeleton  of  shank 

2.304 

763.4 

558.2 

68.4 

486.2 

86.63 

Skeleton  of  flank  and  plate 

Skeleton  of  rump 

1,862 

622 

990.4 

219.0 

225.0 

127.2 

62.9 

18.9 

225.1 

201.9 

54.04 

26.25 

Skeleton  of  chuck  and  neck 

4.896 

2021.3 

721.7 

159.8 

962.8 

172.93 

Skeleton  of  round 

2.350 

829.2 

643.2 

59.3 

392.6 

65.82 

Skeleton  of  loin 

2,550 

887.6 

698.2 

71.0 

489.8 

91.16 

Skeleton  of  rib 

1,824 

823.3 

342.4 

60.1 

417.0 

74.58 

Horns 

304 

165.0 

1.8 

16.6 

37.9 

7.33 

Hoofs  and  dewclaws* 

481 

320.5 

2.2 

25.8 

4.9 

0.32 

Teethf 

278 

79.7 

3.1 

5.5 

150.6 

28.36 

*Hoofs  and  dewclaws  of  steer  538  and  steer  540  were  analyzed  together. 
fTeeth  of  steer  538  and  steer  540  were  analyzed  together. 


Studies  In  Animal  Nutrition — III 


67 


Table  51. — Steer  541.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

12,470 

10239.4 

11.1 

345.8 

84.7 

3.37 

Circulatory  system 

2,646 

1282.3 

1055.2 

43.6 

15.1 

2.78 

Respiratory  system 

2,387 

1890.9 

47.2 

65.4 

24.8 

5.08 

Brain  and  spinal  cord 

568 

411.6 

77.3 

9.4 

8.7 

2.11 

Digestive  and  excretory  system  (partial) 

16,313 

11628.7 

2398.0 

330.2 

121.7 

21.86 

Offal  fat 

11,009 

1289.6 

9511.6 

32.3 

13.8 

2.86 

Heart  and  neck  sweetbreads 

740 

484.3 

141.7 

16.9 

10.8 

2.72 

Liver 

3,832 

2664.8 

95.9 

121.0 

55.2 

13.37 

Gall 

202 

179.8 

0.4 

0.6 

2.4 

0.12 

Spleen 

596 

463.0 

10.8 

17.1 

7.9 

1.66 

Pancreas 

390 

233.7 

91.4 

7.9 

4.7 

1.11 

Kidneys 

645 

473.9 

63.9 

14.3 

6.9 

1.38 

Hair  and  hide 

26,574 

16057.3 

970.0 

1512.6 

292.3 

14.62 

Head  and  tail,  lean  and  fat 

2,062 

1370.6 

299.4 

60.6 

21.0 

3.38 

Shin  and  shank,  lean  and  fat 

6.830 

4693.9 

616.1 

199.6 

65.3 

12.16 

Flank  and  plate,  lean  and  fat 

24,910 

12405.7 

8529.4 

604.6 

178.6 

31.64 

Rump,  lean  and  fat 

4.454 

2250.7 

1459.8 

105.6 

31.8 

6.10 

Chuck  and  neck,  lean  and  fat 

40,480 

26418.5 

5979.7 

1190.9 

369.6 

66.79 

Round,  lean 

27,000 

19957.1 

908.8 

901.8 

294.0 

56.16 

Round,  fat 

3,854 

817.1 

2828.8 

38.0 

10.9 

1.77 

Loin,  lean 

23,416 

16848.3 

1301.7 

757.0 

245.9 

46.36 

Loin,  fat 

8,088 

1202.0 

6521.7 

63.1 

16.5 

3.15 

Rib,  lean 

11,588 

7956.0 

1209.0 

362.2 

113.1 

21.67 

Rib,  fat 

2,434 

432.9 

1850.8 

26.3 

7.6 

1.44 

Kidney, fat 

6,056 

272.2 

5713.8 

12.2 

5.2 

1.21 

Skeleton  of  feet 

4,506 

1872.3 

578.6 

154.9 

973.7 

168.66 

Skeleton  of  head 

5,400 

2730.2 

381.3 

161.7 

1085.8 

197.48 

Skeleton  of  tail 

206 

98.7 

40.4 

6.5 

22.4 

3.90 

Skeleton  of  shin 

2,946 

851.4 

690.0 

91.4 

799.5 

111.89 

Skeleton  of  shank 

3,744 

1222.7 

778.5 

109.8 

1039.5 

141.45 

Skeleton  of  flank  and  plate 

2.864 

1487.0 

439.0 

65.2 

339.1 

51.52 

Skeleton  of  rump 

1,056 

319.9 

258.3 

32.3 

268.7 

51.25 

Skeleton  of  chuck  and  neck 

7,296 

2542.6 

1341.7 

245.7 

1832.0 

264.26 

Skeleton  of  round 

3,250 

876.1 

1054.9 

105.7 

631.5 

136.86 

Skeleton  of  loin 

3,916 

1344.9 

968.6 

125.9 

786.4 

152.92 

Skeleton  of  rib 

3,162 

1001.5 

682.3 

101.0 

807.5 

117.82 

Horns 

468 

248.3 

3.1 

25.0 

64.8 

12.59 

Hoofs  and  dewclaws 

869 

505.1 

5.2 

52.3 

10.9 

0.53 

Teeth 

304 

142.4 

2.9 

4.4 

123.8 

23.79 

Table  52. — Steer  547.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

8,711 

7042.8 

256.3 

62.5 

2.53 

Circulatory  system 

1,624 

949.1 

451.7 

31.9 

11.1 

2.01 

Respiratory  system 

1,868 

1474.0 

52.6 

47.6 

21.5 

3.64 

Brain  and  spinal  cord 

459 

354.2 

41.1 

7.1 

6.7 

1.68 

Digestive  and  excretory  system  (partial) 

11,978 

8883.2 

1309.1 

256.1 

124.8 

23.00 

Offal  fat 

3,879 

700.4 

3059.8 

23.5 

10.0 

2.06 

Liver 

2,851 

2051.7 

73.6 

84.3 

38.9 

10.43 

Spleen 

391 

301.4 

8.7 

11.8 

5.3 

1.17 

Pancreas 

200 

142.8 

20.2 

4.9 

2.9 

0.75 

Kidneys 

450 

329.7 

38.5 

11.5 

5.2 

1.08 

Hair  and  hide 

14,618 

9435.3 

267.4 

779.1 

189.3 

10.52 

Head,  tail,  shin  and  shank,  lean  and  fat 

8,590 

5906.7 

1053.2 

242.1 

77.1 

14.86 

Flank  and  plate,  lean  and  fat 

14,226 

7829.0 

3915.4 

355.9 

112.8 

20.91 

Rump,  lean  and  fat 

2,256 

1227.5 

622.1 

55.6 

18.5 

3.54 

Chuck  and  neck,  lean  and  fat 

23,636 

15975.1 

2954.7 

655.2 

217.0 

39.47 

Round,  lean 

17,092 

12723.3 

612.6 

550.9 

184.4 

35.89 

Round,  fat 

2,150 

642.3 

1304.3 

28.9 

8.5 

1.46 

Loin,  lean 

13,576 

9708.5 

831.4 

432.0 

139.8 

27.02 

Loin,  fat 

3,972 

837.4 

2869.7 

39.2 

11.9 

2.26 

Rib,  lean 

6,560 

4579.0 

618.8 

199  6 

64.2 

1.18 

Rib,  fat 

1,102 

276.4 

730.5 

15.9 

5.5 

0.83 

Kidney,  fat 

1,630 

128.6 

1468.7 

5.1 

2.2 

0.24 

Skeleton  of  feet,  head,  tail,  shin  and  shank.. 

11,966 

5383.9 

1685.5 

399.8 

2272.2 

418.93 

Skeleton  of  flank  and  plate 

1.958 

1044.4 

262.8 

60.9 

230.5 

42.45 

Skeleton  of  rump 

760 

:w,  o 

121.0 

25.0 

180.0 

33.77 

Skeleton  of  chuck  and  neck 

5,120 

2252.3 

715.9 

164  9 

1035.6 

186.62 

Skeleton  of  round 

2,488 

902  6 

711.8 

62.8 

430.8 

81.06 

Skeleton  of  loin 

3,132 

1233.5 

629.5 

91.1 

631.3 

120.30 

Skeleton  of  rib 

2,172 

902.1 

368.2 

72.1 

423.0 

78.89 

Horns,  hoofs  and  dewciaws 

737 

392.2 

9.8 

53.9 

18.2 

1.62 

Teeth 

310 

100.8 

2.1 

6.8 

158.7 

29.61 

68 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  53. — Steer  548.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

4,603 

3737.5 

142.7 

32.9 

0 97 

Circulatory  system 

753 

528.5 

94.7 

19.6 

6.4 

1.18 

Respiratory  system 

1,084 

833.4 

33.9 

31.5 

2.0 

2.19 

Brain  and  spinal  cord 

526 

392.6 

58.2 

8.6 

7.5 

1.84 

Digestive  and  excretory  system  (partial) 

5,357 

4230.5 

228.6 

127.8 

7.1 

9.59 

Offal  fat 

845 

444.2 

310.8 

12.4 

6.1 

1.03 

Heart  and  neck  sweetbreads 

168 

129.5 

10.2 

4.7 

2.7 

0.66 

Liver 

1,104 

781.8 

20.3 

36.3 

15.3 

3.71 

Spieen 

215 

166.1 

2.9 

6.7 

2.9 

0.60 

Pancreas 

85 

64.1 

5.3 

2.4 

1.2 

0.28 

Kidneys 

353 

277.3 

12.0 

8.8 

4.4 

0.87 

Hair  and  hide 

8,358 

5466.8 

78.0 

466.8 

112.8 

6.52 

Head  and  tail,  lean  and  fat 

967 

693.2 

87.0 

25.9 

9.2 

1.69 

Shin  and  shank,  lean  and  fat 

2,462 

1842.7 

109.5 

80.4 

24.6 

4.31 

Flank  and  plate,  lean  and  fat 

4,802 

3379.8 

366.3 

159.4 

46.5 

8.36 

Rump,  lean  and  fat 

1,042 

756.3 

60.4 

34.9 

11.5 

2.13 

Chuck  and  neck,  lean  and  fat 

11,136 

8421.6 

390.4 

345.2 

117.3 

20.82 

Round,  lean 

9,208 

6953.3 

154.7 

309.7 

103.0 

19.71 

Round,  fat 

520 

279.9 

149.4 

14.2 

3.6 

0.45 

Loin,  lean 

5,668 

4269.4 

114.5 

186.6 

63.9 

11.85 

Loin,  fat 

384 

157.2 

175.0 

7.6 

2.3 

0.37 

Rib,  lean  and  fat 

3,180 

2412.0 

87.1 

99.6 

34.8 

6.20 

Kidney,  fat 

284 

78.5 

182.4 

3.9 

1.4 

0.17 

Skeleton  of  feet 

2,496 

1138.7 

345.1 

83.5 

459.6 

80.25 

Skeleton  of  head 

2,989 

1702.2 

202.8 

82.8 

511.4 

83.65 

Skeleton  of  tail 

94 

53.3 

10.3 

3.2 

10.0 

1.82 

Skeleton  of  shin 

1,584 

617.4 

312.2 

51.0 

308.7 

56.52 

Skeleton  of  shank 

1,712 

615.7 

362.9 

61.9 

319.8 

59  34 

Skeleton  of  flank  and  plate 

1,514 

875.8 

163.2 

48.4 

136.0 

23.41 

Skeleton  of  rump 

570 

250.2 

90.8 

18.5 

104.2 

18.94 

Skeleton  of  chuck  and  neck 

3,630 

1718.4 

479.2 

112.9 

623.5 

116.45 

Skeleton  of  round 

1,898 

767.2 

496.8 

49.7 

311.4 

57.26 

Skeleton  of  loin 

1,652 

726.8 

312.9 

51.7 

281.3 

53.36 

Skeleton  of  rib 

1,614 

753.3 

237.1 

55.2 

265.7 

48.50 

Horns,  hoofs  and  dewclaws 

435 

241.8 

5.0 

30.1 

11.3 

1.15 

Teeth 

264 

111.2 

1.7 

5.5 

111.3 

20.95 

Table  54. — Steer  550.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood  

7,080 

5760.4 

203.0 

43.1 

2.19 

Circulatory  system 

1,163 

651.3 

347.9 

23.1 

7.7 

1.55 

Respiratory  system 

1,460 

1137.0 

58.2 

36.2 

14.6 

2.91 

Brain  and  spinal  cord 

443 

333.1 

44.7 

7.0 

6.4 

1.55 

Digestive  and  excretory  system  (partial) .... 

9,156 

6653.2 

1192.0 

179.0 

107.7 

16.94 

Offal  fat 

2,610 

544.9 

1949.3 

15.3 

8.4 

2.06 

Liver 

1,992 

1398.4 

62.5 

59.6 

26.4 

6.22 

Spleen 

291 

233.4 

7.7 

8.6 

4.3 

0.95 

Pancreas 

200 

133.8 

31.1 

4.9 

2.7 

0.64 

Kidneys 

379 

285.6 

23.7 

9.5 

4.5 

0.96 

Hair  and  nide 

10,440 

6779.2 

127.1 

555.4 

136.5 

8.77 

Head,  tail,  shin,  and  shank,  lean  and  fat 

5,274 

3752.2 

491.6 

149.2 

49.9 

9.18 

Flank  and  plate,  lean  and  fat 

7,896 

4868.1 

1532.3 

225.6 

70.1 

12.95 

Rump,  lean  and  fat 

1,662 

995.3 

344.8 

44.7 

15.3 

2.96 

Chuck  and  neck,  lean  and  fat 

15,814 

10633.2 

2229.6 

424.6 

150.2 

23.47 

Round,  lean 

11,720 

8849.2 

310.9 

400.6 

124.6 

24.96 

Round,  fat 

944 

306.2 

546.8 

13.6 

3.9 

0.65 

Loin,  lean 

9,072 

6709.2 

408.5 

281.0 

96.1 

20.96 

Loin,  fat 

2,134 

465.6 

1524.7 

19.6 

7.0 

1.39 

Rib,  lean  and  fat 

4,408 

3045.1 

486.4 

128.9 

43.9 

8.24 

Kidney,  fat 

756 

78.9 

656.8 

3.4 

1.7 

0.43 

Skeleton  of  feet,  head,  tail,  shin  and  shank. . . 

9,839 

4520.0 

1539.4 

290.2 

1836.8 

340.53 

Skeleton  of  flank  and  plate 

1,540 

810.0 

197.0 

49.9 

187.3 

33.37 

Skeleton  of  rump 

700 

285.6 

133.7 

23.7 

133.1 

25.45 

Skeleton  of  chuck  and  neck 

4,956 

2224.1 

810.9 

153.5 

895.2 

163.25 

Skeleton  of  round 

2,100 

776.9 

602.5 

55.2 

355.1 

67.35 

Skeleton  of  loin 

2,464 

948.2 

610.0 

71.4 

438.0 

82.30 

Skeleton  of  rib 

1,740 

730.6 

306.7 

57.4 

322.1 

59.23 

Homs,  hoofs  and  dewclaws 

549 

292.1 

7.3 

40.1 

13.5 

1.21 

Teeth 

228 

74.1 

1.6 

5.0 

116.7 

21.77 

Studies  In  Animal  Nutrition — III 


69 


Table  55. — Steer  552.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

5,219 

4163.4 

157.1 

44.6 

1.46 

1,056 

1.096 

466 

676.6 

219.5 

22.9 

8.3 

1 37 

853.8 

28.7 

29.9 

13.2 

2.25 

345.2 

53.5 

7.3 

6 4 

1 56 

Digestive  and  excretory  system  (partial) .... 

5.936 

1.784 

4452.5 
516  0 

566.2 

1208.8 

128.2 

16.5 

62.8 

6.8 

10.33 
1 23 

265 

166.0 

62.3 

6.2 

3 8 

0 95 

1.181 

821.0 

24.4 

36.9 

15.7 

3.98 

284 

221.1 

3.7 

8.8 

3 8 

0 79 

101 

74  0 

9.4 

2.5 

1 3 

0 30 

316 

230.3 

38.2 

7 3 

3.6 

0 70 

10,532 

7037.8 

161.6 

508.5 

148.0 

7.37 

Head  and  tail,  lean  and  fat 

1,209 

2.976 

822  2 

175.1 

29  9 

11.3 

1.87 

5.12 

Shin  and  shank,  lean  and  fat 

2216.2 

138.0 

95.6 

29.1 

Flank  and  plate,  lean  and  fat 

6,204 

3928.9 

1088.0 

177.8 

54.7 

9.00 

Rump,  lean  and  fat 

1.326 

869.6 

196.3 

38.3 

12.9 

2.32 

Chuck  and  neck,  lean  and  fat 

12.684 

9248.5 

899.0 

378.9 

126.1 

22.58 

Round,  lean 

9.830 

7479.7 

192.4 

320.0 

111.0 

20.35 

Round,  fat 

908 

404.8 

378.3 

19.2 

5.4 

0 61 

Loin,  lean 

7,034 

1.042 

5244.5 

25.2 

220.2 

77.4 

13.65 

Loin,  fat 

233.8 

739.7 

12.3 

4.1 

0 64 

Rib.  lean  and  fat 

4,104 

2971.5 

264.1 

132.7 

39.8 

7.39 

Kidnev  fat 

500 

64.6 

422.8 

2 7 

1.4 

0 17 

Skeleton  of  feet 

2,861 

3.052 

1243.2 

366.9 

99.5 

559.8 

99.56 

Skeleton  of  head 

1652.4 

190.3 

89.0 

574.1 

105.39 

Skeleton  of  tail 

131 

70  6 

17.5 

3 8 

16.8 

3.04 

Skeleton  of  shin 

1,550 

617.2 

273.8 

55.1 

291.3 

51.65 

Skeleton  of  shank 

1.742 

566.8 

363.1 

53.8 

434.1 

79.84 

Skeleton  of  flank  and  plate 

1,516 

640 

822.8 

192.8 

51.1 

161.5 

28.97 

Skeleton  of  rump 

254.7 

110.4 

21.1 

132.8 

23.49 

Skeleton  of  chuck  and  neck 

3,908 

1,626 

2,192 

1749.0 

588.2 

126.0 

718.3 

130.68 

Skeleton  of  round 

541.6 

482.5 

41.7 

322.1 

56.65 

Skeleton  of  loin 

899.9 

458.4 

63.0 

402.8 

75  78 

Skeleton  of  rib 

2,084 

550 

912.8 

337.4 

68.4 

365  0 

68.90 

Horns,  hoofs  and  dev  claws 

317  3 

10.1 

35.3 

14.8 

1 44 

Teeth 

278 

113.8 

2.2 

5.7 

120.2 

22.83 

Table  56. — Steer  554.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood  

4.197 

3402  6 

121.3 

33.6 

1.30 

Circulatory  system 

591 

428.9 

61.5 

5.0 

5.4 

1.00 

Respiratory  system 

907 

716.7 

22.1 

24.4 

11.0 

2.06 

Brain  and  spinal  cord 

395 

303.5 

34.7 

7.0 

6.2 

1.38 

Digestive  and  excretory  system  (partial) .... 
Offal  fat 

4.216 

644 

3276.5 

265.6 

324.9 

327.6 

89.9 

7.7 

46.3 

4.1 

7.97 

0.64 

Heart  and  neck  sweetbreads 

336 

271.1 

7.2 

8.9 

7.1 

1.59 

Liver 

1,166 

824  8 

30.2 

32.8 

19.7 

4.00 

Spleen 

202 

158.2 

2.9 

6.0 

3.0 

0.63 

Pancreas 

76 

53.5 

10.1 

1.8 

1.1 

0.21 

Kidneys 

530 

432.0 

12.7 

11.8 

6.1 

1.14 

Hair  and  hide 

7,400 

883 

4885.3 

119.5 

368.2 

106.7 

7.10 

Head  and  tail,  lean  and  fat 

642.6 

73.7 

24.8 

11.1 

1.69 

Shin  and  shank,  lean  and  fat 

2,304 

4,232 

1.052 

1715.4 

90.9 

76.5 

25.9 

4.06 

Flank  and  plate,  lean  and  fat 

2993.7 

376.5 

126.8 

41.5 

7.19 

Rump,  lean  and  fat 

741.1 

95.6 

32.2 

12.3 

1.87 

Chuck  and  neck,  lean  and  fat 

10,530 

7,918 

520 

7882.0 

465.0 

326.6 

131.1 

19.90 

Round,  lean 

6013  7 

171.3 

260.2 

100.6 

17.02 

Round,  fat 

267  5 

169.9 

13.9 

4.0 

0.43 

Loin,  lean 

5,812 

428 

4350.1 

215.0 

185.9 

66.9 

12.03 

Loin,  fat 

127.3 

247.7 

6.0 

2.3 

0.34 

Rib,  lean  and  fat 

2,914 

240 

2179.5 

100.8 

95.0 

33.2 

5.77 

Kidney, fat 

44.6 

181.0 

1.7 

0.8 

0.14 

Skeleton  of  feet 

2,403 

2,229 

1117.3 

335.9 

89.0 

425.0 

78.91 

Skeleton  of  head 

1330.9 

70.4 

69.3 

375.6 

70.26 

Skeleton  of  tail 

125 

71 .0 

14.2 

4.3 

13.8 

2.58 

Skeleton  of  shin 

1.464 

585.7 

269.6 

47.2 

307.2 

58.30 

Skeleton  of  ehank 

1.918 

772.3 

371.3 

62.3 

334.8 

59.92 

Skeleton  of  flank  and  plate 

1.278 

775.5 

127.2 

42.0 

99.1 

17.56 

Skeleton  of  rump 

Skeleton  of  chuck  and  neck 

732 

3,732 

1,702 

357.2 

1920.5 

92.7 

374.5 

25.4 

117.7 

119.2 

611.6 

21.59 

109.91 

Skeleton  of  round 

684.2 

420  4 

45.4 

271.5 

46.07 

Skeleton  of  loin 

1.868 

963.9 

258.7 

55.9 

280.2 

50.25 

Skeleton  of  rib 

1.354 

690.3 

166.7 

43.0 

207.7 

36.02 

Horns,  hoofs  and  dewclaws 

338 

186.8 

4.9 

25.9 

8.1 

0.77 

Teeth 

225 

104.2 

0.2 

6.6 

87.1 

14.31 

70 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  57. — Steer  555.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

4,529 

3626.6 

141.5 

33.4 

1 59 

Circulatory  system 

554 

393.2 

66.8 

14.3 

4.9 

0.88 

Respiratory  system 

937 

744.4 

26.0 

25.2 

10.5 

1.95 

Brain  and  spinal  cord 

353 

269.6 

35.1 

6.0 

5.4 

1.25 

Digestive  and  excretory  system  (partial) .... 

4,534 

3553.6 

288.2 

105.1 

49.5 

8.57 

Offal  fat 

462 

271.2 

144.9 

8.0 

3.6 

0.55 

Heart  and  neck  sweetbreads 

159 

120.8 

10.1 

4.3 

2.1 

0.44 

Liver 

1.240 

925.5 

31.5 

36.4 

18.5 

4.33 

Spleen 

167 

129.2 

2.3 

5.2 

2.2 

0.46 

Pancreas 

106 

80.8 

5.8 

2.8 

1.6 

0.31 

Kidneys 

439 

360.0 

12.0 

9.8 

5.2 

0.98 

Hair  and  hide 

6,580 

4543.6 

49.7 

342.2 

97.1 

5 86 

Head  and  tail,  lean  and  fat 

956 

694.8 

88.3 

25.1 

10.1 

1.55 

Shin  and  shank,  lean  and  fat 

2,688 

2071.0 

62.7 

85.7 

28.1 

4.87 

Flank  and  plate,  lean  and  fat 

4,068 

3071.9 

139.3 

129.3 

42.2 

7.12 

Rump,  lean  and  fat 

876 

630.5 

63.7 

25.5 

9.4 

1.64 

Chuck  and  neck,  lean  and  fat 

9,402 

7281.9 

201.5 

294.3 

97.9 

17.39 

Round,  lean 

7,126 

5554.7 

96.0 

218.0 

85.1 

14.75 

Round,  fat 

376 

221.8 

89.5 

9.8 

2.8 

0.34 

Loin,  lean 

4,618 

3599.2 

79.7 

143.4 

55.4 

9.60 

Loin,  fat 

274 

121.0 

117.7 

6.0 

2.3 

0.27 

Rib,  lean  and  fat 

2,512 

1949.7 

63.7 

77.9 

29.5 

4.75 

Kidney,  fat 

130 

42.9 

74.6 

1.6 

0.7 

0.12 

Skeleton  of  feet 

2,157 

1110.0 

195.0 

87.2 

350.5 

59.99 

Skeleton  of  head 

1,978 

1235.0 

63.6 

56.6 

293.0 

50.83 

Skeleton  of  tail 

74 

46.9 

4.9 

2.7 

6.8 

1.13 

Skeleton  of  shin 

1,454 

731.9 

160.9 

47.0 

259.2 

46.38 

Skeleton  of  shank 

1,704 

846.9 

235.3 

49.1 

253.3 

45.26 

Skeleton  of  flank  and  plate 

1,236 

805.5 

70.3 

41.6 

99.3 

16.82 

Skeleton  of  rump 

506 

288.9 

29.9 

17.7 

81.7 

13.99 

Skeleton  of  chuck  and  neck 

3,216 

1907.4 

165.7 

101.1 

447.4 

79.21 

Skeleton  of  round 

1,652 

884.9 

219.0 

46.1 

244.5 

42.65 

Skeleton  of  loin 

1,264 

749.6 

90.6 

39.2 

164.4 

29.24 

Skeleton  of  rib 

1,256 

734.7 

79.7 

41.1 

165.7 

28.54 

Horns,  hoofs  and  dewciaws 

298 

145.6 

2.7 

22.6 

7.9 

0.40 

Teeth 

190 

81.0 

0.6 

5.5 

81.4 

13.86 

Table  58. — Steer  556.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

6,124 

4989.4 

170.1 

48.0 

2.14 

Circulatory  system 

993 

619.2 

237.9 

21.3 

8.5 

1.47 

Respiratory  system 

1,272 

988.1 

43.5 

34.3 

17.8 

2.75 

Brain  and  spinal  cord 

398 

267.1 

77.4 

6.5 

6.4 

1.39 

Digestive  and  excretory  system  (partial) 

6,003 

4473.6 

631.0 

132.3 

78.3 

14.47 

Offal  fat 

1,402 

457.3 

859.6 

13.5 

6.3 

0.88 

Heart  and  neck  sweetbreads 

328 

250.6 

21.6 

8.5 

67.3 

1.48 

Liver 

1,760 

1251.6 

61.6 

53.9 

40.4 

6.44 

Spleen 

300 

233.2 

7.0 

9.0 

4.4 

0.82 

Pancreas 

96 

69.4 

10.2 

2.3 

1.4 

0.29 

Kidneys 

338 

254.6 

24.0 

8.5 

4.5 

0.86 

Hair  and  hide 

10,314 

6836.5 

222.1 

531.9 

142.0 

9.08 

Head  and  tail . lean  and  fat 

914 

628.3 

120.6 

25.1 

8.2 

1.51 

Shin  and  shank,  lean  and  fat 

2,808 

2070.4 

126.6 

91.9 

27.1 

5.19 

Flank  and  plate,  lean  and  fat 

6,174 

4153.1 

824.8 

194.1 

57.7 

10.37 

Rump,  lean  and  fat 

1,244 

862.5 

129.2 

37.5 

12.9 

2.38 

Chuck  and  neck,  lean  and  fat 

12,406 

8971.5 

944.7 

374.2 

154.6 

22.33 

Round,  lean 

9,472 

7121.0 

302.6 

311.5 

118.5 

19.89 

Round,  fat 

686 

305.7 

298.7 

15.1 

4.0 

0.47 

Loin,  lean 

6,938 

5147.0 

262.2 

226.9 

82.5 

14.29 

Loin,  fat 

820 

214.1 

541.5 

11.5 

4.4 

0.52 

Rib,  lean  and  fat 

3,300 

2357.5 

238.4 

105.1 

38.1 

6.07 

Kidney,  fat 

420 

64.5 

338.6 

2.5 

1.5 

0.16 

Skeleton  of  feet 

2,589 

1202.4 

367.0 

96.5 

459.8 

84.89 

Skeleton  of  head 

2,610 

1478.0 

187.6 

78.0 

460.1 

85.37 

Skeleton  of  tail 

92 

52.4 

9.2 

3.3 

12.3 

2.20 

Skeleton  of  shin 

1,702 

685.5 

337.0 

55.8 

336.6 

62.89 

Skeleton  of  shank 

1,952 

731.1 

441.4 

66.2 

389.8 

73.92 

Skeleton  of  flank  and  plate 

1,578 

909.9 

18.4 

52.3 

162.0 

29.38 

Skeleton  of  rump 

608 

286.9 

80.1 

21.5 

110.4 

20.72 

Skeleton  of  chuck  and  neck 

3,734 

1758.9 

468.1 

130.2 

692.5 

132.37 

Skeleton  of  round 

1,974 

753.0 

507.8 

52.6 

329.7 

63.40 

Skeleton  of  loin 

2,268 

1046.8 

361.9 

71.6 

384.0 

75.05 

Skeleton  of  rib 

1,646 

763.2 

203.3 

54.5 

298.9 

61.23 

Horns,  hoofs  and  dewciaws 

390 

187.9 

4.8 

32.4 

7.3 

0.60 

Teeth 

218 

91.0 

0.1 

6.4 

95.8 

15.06 

Studies  In  Animal  Nutrition — III 


71 


Table  59. — Steer  557.  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phospnorus 

8,952 

2.342 

7156.3 

280.7 

75.4 

2.42 

1183.1 

903.0 

38.6 

14.4 

2.46 

1.972 

1519.5 

103.7 

51.2 

22.1 

4.00 

399.9 

68.5 

69.9 

7.8 

1 86 

Digestive  and  excretory  system  (partial) 

9,981 

6,757 

623 

7371.1 

942.3 

1142.4 

5640.1 

200.3 

29.4 

104.9 

15.5 

17.57 
1 .96 

422.3 

105.2 

14.5 

10.3 

2 22 

3,003 

485 

2086.0 

56.5 

92.6 

40.4 

9.97 

373.1 

16.2 

14.0 

6.7 

1.36 

225 

145.9 

38.6 

5.1 

2.4 

0.56 

649 

498.1 

38.2 

15.4 

7.5 

1.41 

14,100 

1,915 

4,196 

8900.5 

706.7 

703.3 

165.1 

10.01 

1158.1 

462.2 

45.0 

16  2 

2.76 

Shin  and  shank,  lean  and  fat 

2861.9 

496.3 

123.1 

38.9 

6.42 

Flank  and  plate,  lean  and  fat 

15,294 

7713.4 

5251.2 

357.0 

119.0 

20.34 

Rump,  lean  and  fat 

2,582 

22,146 

14,960 

2,532 

11,714 

4,472 

6,372 

1,616 

3,288 

3,558 

3,969 

193 

1321.4 

848.1 

59.8 

20.6 

3.67 

Chuck  and  neck,  lean  and  fat 

14152.0 

3891.9 

598.4 

211.9 

40  53 

Round,  lean 

11044.1 

689.7 

459.7 

151.3 

29.02 

Round,  fat 

702.5 

1611.5 

25.8 

8.8 

1 22 

Loin,  lean 

8335.1 

916.7 

347.4 

128.6 

22.49 

Loin,  fat  

711.4 

3573.1 

32.6 

11.1 

1 70 

Rib,  lean  

4322.2 

763.8 

185.1 

61.9 

10.83 

0.95 

Rib,  fat 

316.3 

1194.5 

15.5 

5.8 

Kidney, fat 

266.9 

2897.7 

11.9 

5.1 

0.77 

Skeleton  of  feet 

1579.2 

426.8 

128.1 

670.9 

124.10 

Skeleton  of  head 

2136.0 

248.6 

109.2 

724.9 

130 . 74 

Skeleton  of  tail 

102.3 

32.4 

6.1 

20.2 

3.77 

Skeleton  of  shin 

2,246 

2,010 

2.498 

960 

826.4 

404.6 

75.3 

477.0 

88.96 

Skeleton  of  shank 

962.7 

510.0 

88.9 

523.4 

94.61 

Skeleton  of  dank  and  plate 

1408.7 

316.2 

78.7 

242.5 

42.99 

Skeleton  of  rump 

410.4 

128.7 

33.0 

183.9 

35.62 

Skeleton  of  chuck  and  neck 

5,638 

2421.1 

680.7 

191.4 

1175.4 

212.21 

Skeleton  of  round 

2,686 

2,478 

2,572 

695 

873.1 

789.6 

69.1 

514.8 

95.30 

Skeleton  of  loin 

957.2 

499.1 

73.3 

480.3 

90.57 

Skeleton  of  rib 

1037.8 

432.9 

83.3 

505.9 

92.87 

Horns,  hoofs  and  dewclaws 

373.0 

9.8 

48.3 

17.9 

19.39 

Teeth 

261 

104.4 

0.7 

7.5 

120.4 

22.53 

Table  60. — Steer  558 A'  Weights  of  Constituents  in  Samples,  Grams. 


Description  of  sample 

Sample 

Water 

Crude  fat 

Nitrogen 

Ash 

Phosphorus 

Blood 

4,666 

3903.8 

117.1 

32.3 

1 82 

Circulatory  system 

971 

643.2 

184.8 

21.6 

7.1 

1.39 

Respiratory  system 

1,111 

884.7 

16.4 

29.8 

11.7 

2.57 

Brain  and  spinal  cord 

520 

393.1 

52.0 

8.3 

7.8 

1.94 

Digestive  and  excretory  system  (partial) .... 

6,510 

5075.3 

414.4 

145.9 

64.6 

12.24 

Offal  fat 

829 

394.6 

358.2 

11.5 

4.2 

0.83 

Liver 

1,352 

966.3 

28.4 

41.9 

18.2 

4.76 

Spleen 

193 

146.3 

2.1 

6.4 

3.0 

0.54 

Pancreas 

153 

116.7 

7.9 

3.9 

2.2 

0.57 

Kidneys 

318 

244.2 

13.8 

8.3 

3.8 

0.79 

Hair  and  hide 

8,138 

5266.3 

87.7 

430.3 

92.0 

6.75 

Head,  tail,  shin,  and  shank,  lean  and  fat.  . . . 

4,324 

3216.2 

238.2 

127.5 

40.8 

7.05 

Flank  and  plate,  lean  and  fat 

4,192 

2961.7 

306.5 

135.4 

38.7 

7.21 

Rump,  lean  and  fat 

1,030 

719.4 

99.3 

31.8 

10.4 

2.02 

Chuck  and  neck,  lean  and  fat 

11,682 

8789.8 

481.4 

355.3 

108.3 

21.85 

Round,  lean 

9,664 

7456.0 

107.6 

301.1 

101.4 

20.58 

Round,  fat 

644 

289.8 

266.2 

14.9 

3.9 

0.61 

Loin,  lean 

5,962 

4468.5 

158.1 

189.7 

61.1 

12.52 

Loin,  fat 

600 

181.2 

356.5 

9.3 

2.7 

0.48 

Rib,  lean  and  fat 

3,242 

2421.1 

113.1 

102.4 

32.1 

6.39 

Kidney  fat 

220 

40.5 

165.4 

2.1 

0.6 

0.12 

Skeleton  of  feet,  head,  tail  .shin  and  shank 

9,785 

4475.4 

1645.3 

303  0 

1716.4 

318.21 

Skeleton  of  flank  and  plate 

1,092 

596.6 

137.0 

36.1 

113.7 

20.58 

Skeleton  of  rump 

678 

288.5 

124.6 

21.1 

126.6 

22.94 

Skeleton  of  chuck  and  neck 

4,056 

1752.7 

733.8 

125.3 

723.9 

131.41 

Skeleton  of  round 

2,094 

717.6 

708.0 

47.9 

338.4 

62.44 

8keleton  of  loin 

2,138 

850.3 

535.3 

54.1 

381.4 

70.15 

Skeleton  of  rib 

1,516 

710.8 

243.3 

50.8 

230.3 

41.64 

Homs,  hoofs  and  dewclaws 

443 

235.7 

5.9 

32.4 

10.9 

0.98 

Teetn 

274 

89.1 

1.9 

6.0 

140.3 

26.17 

72 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  61. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 

(3  Months-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorus 

Animal  556,  Group  1 

Lean  and  fat  flesh 

70.59 

9.14 

3.09 

1.13 

0.184 

Bone 

46.59 

14.37 

3.29 

17.52 

3.332 

81.47 

2.78 

0.78 

0 035 

Circulatory  system 

62.36 

23.96 

2.15 

0.86 

0.148 

Respiratory  system 

77.68 

3.42 

2.70 

1.40 

0.216 

Nervous  system 

67.12 

19.45 

1.62 

1.62 

0.349 

Digestive  and  excretory  system 

74.03 

8.56 

2.43 

1.54 

0.276 

Hair  and  Hide 

66.28 

2.15 

5.16 

1.38 

0.088 

Offal  fat 

32.62 

61.31 

0.96 

0.45 

0.063 

Total  animal 

65.23 

9.71 

3.24 

4.88 

0.868 

Animal  554,  Group  II 

Lean  and  fat  flesh 

73.19 

5.94 

3.12 

1.17 

0.191 

Bone 

49.29 

13.30 

3.20 

16.20 

2.932 

Blood 

81.07 

2.89 

0.80 

0.031 

Circulatory  system 

72.58 

10.40 

2.53 

0.91 

0.169 

Respiratory  system 

79.02 

2.44 

2.69 

1.21 

0.227 

Nervous  system 

76.83 

8.78 

1.77 

1.58 

0.350 

Digestive  and  excretory  system 

76.86 

5.94 

2.32 

1.28 

0.238 

Hair  and  hide 

66.02 

1.62 

4.98 

1.44 

0.096 

Offal  fat 

41.23 

50.87 

1.20 

0.63 

0.099 

Total  animal 

67.05 

7.35 

3.23 

4.97 

0.866 

Animal  555,  Group  III 

Lean  and  fat  flesh 

76.42 

3.26 

3.08 

1.10 

0.189 

Bone 

56.63 

7.97 

3.21 

14.34 

2.510 

Blood 

80.08 

3.12 

0.74 

0.035 

Circulatory  system 

70.98 

12.06 

2.57 

0.88 

0.159 

Respiratory  system 

79.45 

2.78 

2.69 

1.12 

0.210 

Nervous  system 

76.37 

9.95 

1.71 

1.54 

0.354 

Digestive  and  excretory  system 

77.80 

5.27 

2.46 

1.19 

0.227 

Hair  and  hide 

69.05 

0.76 

5.20 

1.48 

0.089 

Offal  fat 

58.69 

31.37 

1.72 

0.78 

0.119 

Total  animal 

71.11 

4.38 

3.25 

4.36 

0.739 

Table  62. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 

(5%-Months-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorous 

Animal  557,  Group  1. 

Lean  and  fat  flesh 

58.12 

24.82 

2.48 

0.86 

0.155 

Bone 

Blood 

43.24 

79.94 

15.20 

3.18 

3.14 

18.77 

0.84 

3.440 

0.027 

Circulatory  system 

50.52 

38.56 

1.65 

0.62 

0.105 

Respiratory  system 

77.05 

5.26 

2.60 

1.12 

0.203 

Nervous  system 

72.58 

12.44 

1.79 

1.41 

0.338 

Digestive  and  excretory  system 

72.81 

9.34 

2.29 

1.15 

0.221 

Hair  and  hide 

63.12 

5.01 

4.99 

1.17 

0.071 

Offal  fat 

13.95 

83.47 

0.44 

0.23 

0.029 

Total  Animal 

56.77 

20.99 

2.75 

4.04 

0.731 

Animal  552,  Group  II. 

Lean  and  fat  flesh 

77.03 

9.45 

2.99 

0.99 

0.175 

Bone 

Blood 

43.80 

79.77 

15.87 

3.16 

3.01 

18.68 

0.85 

3.399 

0.028 

Circulatory  system 

64.07 

20.78 

2.17 

0.79 

0.130 

Respitatory  system 

77.90 

2.61 

2.72 

1.21 

0.205 

Nervous  system 

74.07 

11.48 

1.56 

1.37 

0.335 

Digestive  and  excretory  system 

73.80 

8.71 

2.35 

1.13 

0.210 

Hair  and  hide 

66.82 

1.53 

4.83 

1.41 

0.070 

Offal  fat 

28.92 

67.76 

0.92 

0.38 

0.069 

Total  animal 

63.97 

10.48 

3.13 

5.00 

0.880 

Animal  548,  Group  III. 

Lean  and  fat  flesh 

73.75 

4.73 

3.20 

1.05 

0.192 

Bone 

Blood 

46.67 

81.20 

15.25 

3.13 

3.10 

16.87 

0.72 

3. 086 
0.021 

Circulatory  system 

70.18 

12.58 

2.61 

0.84 

0.157 

Respiratory  system 

76.88 

3.13 

2.91 

1.11 

0.202 

Nervous  system 

74.63 

11.06 

1.64 

1.43 

0.350 

Digestive  and  excretory  system 

77.58 

3.84 

2.56 

1.15 

0.216 

Hair  and  hide 

65.41 

0.93 

5.35 

1.35 

0.078 

Offal  fat 

52.57 

36.78 

1.46 

0.72 

0.122 

Total  animal 

66.86 

6.88 

3.32 

4.95 

0.882 

Studies  In  Animal  Nutrition — III 


73 


Table  63. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 


(8%-Months-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorous 

Animal  547,  Group  1. 

Lean  and  fat  flesh 

63.12 

17.92 

2.72 

0.89 

0.156 

Bone 

43.50 

16.29 

3.18 

18.86 

3.486 

80.85 

2.94 

0.72 

0.029 

Circulatory  system 

58.44 

27.81 

1.97 

0.68 

0.124 

Respiratory  system 

78.91 

2.82 

2.55 

1.15 

0.195 

Nervous  system 

77.16 

8.96 

1.54 

1.46 

0.367 

Digestive  and  excretory  system 

73.78 

9.14 

2.32 

1.12 

0.230 

Hair  and  hide 

64.55 

1.83 

5.33 

1.30 

0.072 

Offal  fat 

18.06 

78.88 

0.61 

0.26 

0.053 

Total  Animal 

61.01 

15.73 

2.95 

3.93 

0.704 

Animal  550,  Group  II. 

Lean  and  fat  flesn 

66.53 

14.30 

2.83 

0.94 

0.185 

Bone 

44.11 

17.91 

3.00 

17.86 

3.306 

Blood  

81.36 

2.87 

0 61 

0 031 

Circulatory  system 

56.00 

29.91 

1.99 

0.66 

0.133 

Respiratory  system 

77.88 

3.98 

2.48 

1.00 

0.199 

Nervous  system 

75.20 

10.10 

1.58 

1.44 

0.351 

Digestive  and  excretory  system 

72.35 

10.96 

2.18 

1.21 

0.214 

Hair  and  hide 

64.94 

1.22 

5.32 

1.31 

0.084 

Offal  fat 

20.88 

74.69 

0.59 

0.32 

0.079 

Total  animal 

62.40 

13.94 

2.97 

4.39 

0.798 

Animal  558,  Group  III. 

Lean  and  fat  flesh 

73.49 

5.52 

3.05 

0.96 

0.190 

Bone 

43.97 

19.32 

2.99 

17.00 

3.125 

Blood  

88.67 

2.51 

0.69 

0 039 

Circulatory  system 

66.25 

19.04 

2.22 

0.73 

0.143 

Respiratory  system 

79.63 

1.47 

2.69 

1.05 

0.231 

Nervous  system 

75.60 

10.00 

1.60 

1.51 

0.374 

Digestive  and  excretory  system 

76.81 

5.47 

2.42 

1.08 

0.222 

Hair  and  hide 

64.71 

1.08 

5.29 

1.13 

0.083 

Offal  fat 

47.60 

43.21 

1.39 

0.51 

0.106 

Total  animal 

65.95 

8.59 

3.14 

5.01 

0.914 

Table  64. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 

(ll-Months-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorus 

Animal  541,  Group  1. 

Lean  and  fat  flesh 

58.71 

23.09 

2.68 

0.84 

0.156 

Bone 

Blood 

37.42 

82.11 

18.81 

3.13 

2.77 

22.39 
0 68 

3.646 
0 027 

Circulatory  svstem 

48.46 

39.88 

1.65 

0.57 

0.105 

Respiratory  system 

79.22 

1.98 

2.74 

1.04 

0.213 

Nervous  system 

72.46 

13.61 

1.66 

1.53 

0.371 

Digestive  and  excretory  system 

70.99 

12.33 

2.24 

0.92 

0.186 

Hair  and  hide 

60.43 

3.65 

5.69 

1.10 

0.055 

Offal  fat 

11.71 

86.40 

0.29 

0.13 

0.026 

Total  animal 

56.23 

21.08 

2.91 

3.86 

0.630 

Animal  538,  Group  II. 

Lean  and  fat  flesh 

67.66 

14.01 

2.90 

0.94 

0.176 

Bone 

Blood 

38.47 

82.40 

18.49 

3.08 

2.74 

21.61 

0.79 

3.622 

0.028 

Circulatory  system 

55.48 

28.70 

1.93 

0.63 

0.114 

Respiratory  svstem 

80.80 

2.06 

2.50 

1.02 

0.199 

Nervous  system 

74.33 

11.75 

1.61 

1.50 

0.348 

Digestive  and  excretory  system 

75.01 

8.65 

2.25 

1.00 

0.190 

Hair  and  hide 

64.34 

1.34 

5.56 

1.06 

0.063 

Offal  fat 

21.18 

76.45 

0.54 

0 27 

0.055 

Total  animal 

61.65 

13.73 

3.08 

4.79 

0.799' 

Animal  540,  Group  III. 

Lean  and  fat  flesh 

67.88 

11.72 

2.97 

0 97 

0.179' 

Bone 

Blood 

40.69 
82  28 

17.48 

3.06 

2.81 

19.90 
0 72 

3.692 

0.026 

0.127 

Circulatory  system 

58.13 

27.73 

2.04 

0 6.8 

Respiratory  system 

79.61 

2.31 

2.59 

1.03 

0.198 

Nervous  system 

73.66 

11.94 

1.62 

1.58 

0.375 

Digestive  and  excretory  Bystem 

74.44 

9.41 

2.15 

0.87 

0.169 

Hair  and  hide 

64.64 

2.35 

5.13 

1.26 

0.067 

Offal  fat 

25.24 

71.06 

0.55 

0.28 

0.047 

Total  animal 

62.93 

12.13 

3 07 

4.76 

0.846 

74 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  65. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 
(ll-Montlis-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorus 

Animal  505,  Group  1. 

Lean  and  fat  flesh 

53.69 

29.68 

2.57 

0.73 

0.148 

Bone 

35.79 

17.56 

3.19 

23.85 

4.403 

Blood 

82.26 

2 73 

0 33 

0.023 

0.119 

Circulatory  system 

52.90 

35.03 

1.88 

0.59 

Respiratory  system 

76.83 

5.48 

2.71 

0.95 

0.202 

Nervous  system 

73.81 

14.74 

1.69 

1.72 

0.411 

Digestive  and  excretory  system 

73.04 

11.03 

2.29 

1.03 

0.223 

Hair  and  hide 

62.14 

5.34 

5.30 

0.70 

0.066 

Offal  fat 

12.41 

85.38 

0.34 

0.12 

0.022 

Total  animal 

53.23 

25.06 

2.79 

4.05 

0.749 

Animal  503,  Group  II. 

Lean  and  fat  flesh 

63.17 

18.39 

2.87 

0.84 

0.163 

Bone 

38.28 

15.06 

3.10 

23.70 

4.378 

Blood 

82.78 

2.71 

0.34 

0.076 

Circulatory  system 

51.90 

35.79 

1.75 

0.56 

0.115 

Respiratory  system 

78.71 

3.16 

2.65 

0.98 

0.207 

Nervous  system 

73.11 

16.11 

1.68 

1.55 

0.394 

Digestive  and  excretory  system 

73.23 

8.86 

2.45 

1.08 

0.230 

Hair  and  hide 

67.77 

2.57 

4.79 

0.98 

0.066 

Offal  fat 

14.64 

81.92 

0.52 

0.18 

0.035 

Total  animal 

59.56 

16.36 

2.98 

4.97 

0.913 

Table  66. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 


(18-Months-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorus 

Animal  532,  Group  1. 

Lean  and  fat  flesh 

53.95 

28.72 

2.57 

0.79 

0.142 

Bone 

33.86 

22.22 

3.34 

22.09 

4.016 

Blood 

80.47 

3.00 

0.65 

0.033 

Circulatory  system 

47.92 

40.26 

1.61 

0.57 

0.105 

Respiratory  system 

77.55 

3.41 

2.64 

1.05 

0.193 

Nervous  system 

72.46 

14.46 

1.66 

1.62 

0.392 

Digestive  and  excretory  system 

71.92 

12.02 

2.11 

0.87 

0.177 

hair  and  hide 

59.56 

6.84 

5.23 

1.03 

0.071 

Offal  fat 

9.55 

88.81 

0.24 

0.12 

0.016 

Total  animal 

51.70 

26.74 

2.75 

3.81 

0.680 

Animal  531,  Group  III. 

Lean  and  fat  flesh 

68.05 

10.03 

3.17 

1.03 

0.189 

Bone 

38.05 

16.48 

3.19 

23.84 

4.268 

Blood 

81.88 

2.84 

0.70 

0.040 

Circulatory  system 

56.72 

29.32 

1.97 

0.68 

0.126 

Respiratory  system 

78.34 

2.82 

2.68 

1.05 

0.204 

Nervous  system 

72.99 

13.50 

1.77 

1.50 

0.363 

Digestive  and  excretory  system 

75.88 

8.25 

2.16 

0.85 

0.169 

Hair  and  hide 

63.67 

0.81 

5.73 

1.14 

0.075 

Offal  fat 

25.93 

70.68 

0.62 

0.29 

0.046 

Total  animal 

62.64 

10.75 

3.26 

5.37 

0.950 

Table  68. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 


(3-Year-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorus 

Animal  515,  Group  1,  34  mos. 

Lean  and  fat  flesh 

42.28 

45.22 

1.89 

0.57 

0.103 

Bone 

32.02 

18.72 

3.14 

25.71 

4.153 

Blood 

79.11 

3.22 

0.59 

0.024 

Circulatory  system 

42.03 

48.53 

1.28 

0.48 

0.086 

Respiratory  system 

76.35 

5.44 

2.70 

0.97 

0.187 

Nervous  system 

70.04 

16.09 

1.65 

1.63 

0.395 

Digestive  and  excretory  system 

66.17 

18.71 

2.02 

0.85 

0.162 

Hair  and  hide 

56.00 

12.55 

4.84 

1.86 

0.059 

Offal  fat 

7.53 

90.29 

0.28 

0.12 

0.015 

Total  animal 

43.68 

37.51 

2.29 

3.87 

0.614 

Animal  507,  Group  II,  34  mos. 

Lean  and  fat  flesh 

60.40 

21.48 

2.72 

0.81 

0.151 

Bone 

32.83 

18.05 

3.40 

25.90 

4.004 

Blood 

78.17 

3.41 

0.67 

0.022 

Circulatory  system 

47.12 

41.27 

1.70 

0.54 

0.104 

Respiratory  system 

77.47 

3.93 

2.73 

0.95 

0.159 

Nervous  system 

70.81 

13.74 

1.50 

1.75 

0.418 

Digestive  and  excretory  system 

70.52 

14.08 

2.10 

0.83 

0.150 

Hair  and  hide 

60.85 

6.21 

5.69 

1.07 

0.049 

Offal  fat 

13.10 

83.96 

0.41 

0.14 

0.029 

Total  animal 

56.10 

19.61 

3.03 

5.07 

0.792 

Studies  In  Animal  Nutrition — III 


75 


Table  67. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 

(2-Year-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorous 

Animal  504,  Group  1,  21  mos. 

Lean  and  fat  flesh 

49.66 

35.13 

2.29 

0.68 

0.128 

Bone 

32.99 

17.28 

3.31 

27.14 

4.993 

78.65 

3.33 

0.39 

0 022 

Circulatory  system 

51.05 

36.05 

1.88 

0.69 

0.116 

Respiratory  system 

67.41 

14.90 

2.88 

0.97 

0.181 

Nervous  system 

69.23 

15.08 

1.79 

1.36 

0.350 

Digestive  and  excretory  system 

72.08 

12.76 

2.04 

0.87 

0.178 

Hair  and  hide 

58.29 

8.07 

5.52 

1.06 

0.043 

Offal  fat 

12.76 

84.82 

0.33 

0.15 

0.023 

Total  animal 

49.76 

29.42 

2.64 

4.02 

0.724 

Animal  523,  Group  II,  26  mos. 

Lean  and  fat  flesh 

65.17 

16.50 

2.83 

0.86 

0.101 

Bone 

36.08 

15.39 

3.35 

25.78 

4.725 

Blood 

80.52 

2.79 

0 66 

0.022 

0.110 

Circulatory  system 

53.05 

35.82 

1.68 

0.60 

Respiratory  system 

78.68 

4.05 

2.61 

0.99 

0.189 

Nervous  system 

68.60 

17.61 

1.62 

1.52 

0.359 

Digestive  and  excretory  system 

75.19 

10.33 

2.19 

0.79 

0.158 

Hair  and  hide 

62.80 

1.15 

5.60 

1.03 

0.051 

Offal  fat 

15.42 

81.28 

0.50 

0.20 

0.030 

Total  animal 

60.37 

15.00 

3.10 

5.29 

0.897 

Animal  525,  Group  III,  26  mos. 

Lean  and  fat  flesh 

68.45 

11.91 

2.97 

0.94 

0.174 

Bone 

35.22 

18.20 

3.34 

23.86 

3.952 

Blood  

80.51 

2.94 

0.66 

0 026 

Circulatory  svstem 

64.09 

24.54 

1.71 

0.72 

0.121 

Respiratory  system 

78.61 

2.63 

2.82 

1.11 

0.193 

Nervous  system 

71.05 

13.93 

1.74 

1.50 

0.370 

Digestive  and  excretory  system 

77.48 

9.14 

2.00 

0.88 

0.156 

Hair  and  hide 

62.40 

0.50 

5.72 

1.33 

0.057 

Offal  fat 

21.03 

70.64 

1.29 

0.29 

0.051 

Total  animal 

62.44 

11.87 

3.23 

5.09 

0.838 

Table  69. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 

(40-Months-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorous 

Animal  527,  Group  1. 

Lean  and  fat  flesh 

37.34 

51.80 

1.63 

0.49 

0.089 

Bone 

30  39 

23.73 

3.08 

25.27 

4.302 

Blood  

78.94 

3.28 

0.76 

0 023 

Circulatory  system 

53.95 

32.80 

1.93 

0.64 

0.114 

Respiratory  system 

73.33 

7.19 

2.72 

1.02 

0.166 

Nervous  system 

69.96 

14.06 

1.58 

1.54 

0.368 

Digestive  and  excretory  system 

66.12 

16.84 

2.27 

0.89 

0.145 

Hair  and  hide 

54.48 

11.86 

5.32 

1.37 

0.050 

Offal  fat 

5.39 

93.20 

0.18 

0.12 

0.013 

Thoracic  fat 

10.30 

87.65 

0.34 

0.13 

0.015 

Total  animal 

38.63 

45.45 

2.02 

3.03 

0.507 

Animal  528,  Group  II. 

Lean  and  fat  flesh 

59.20 

22.38 

2.70 

0.83 

0.152 

Bone 

34.14 

19.67 

3.17 

24.10 

4.425 

Blood 

79.93 

3.09 

0.78 

0.025 

Circulatory  system 

63.32 

21.43 

2.34 

0.81 

0.145 

Respiratory  system 

76.62 

3.29 

2.80 

1.03 

0.156 

Nervous  system 

73.22 

10.26 

1.75 

1.73 

0.420 

Digestive  and  excretory  system 

72.16 

10  98 

2.30 

0.92 

0.161 

Hair  and  hide 

57.83 

5.71 

5.92 

1.44 

0.050 

Offal  fat 

12.99 

84.14 

0 41 

0.18 

0.018 

Thoracic  fat 

21.47 

73.14 

0 75 

0 29 

0.034 

Total  animal 

55.33 

20.24 

3.04 

4.92 

0.877 

Animal  524,  Group  III. 

Lean  and  fat  flesh 

70.33 

8.80 

3.15 

0 98 

0.175 

Bone 

37.33 

18.10 

3.10 

23  37 

4.175 

Blood 

82.01 

2 79 

0 75 

0 022 

Circulatory  system 

62  09 

22.42 

2.24 

0.77 

0 132 

Respiratory  system 

77  64 

2 60 

2.79 

1 07 

0.178 

Nervous  system 

73.22 

10.26 

1.60 

1.42 

0.338 

Digestive  and  excretory  system 

73.86 

8.57 

2.41 

1.00 

0.180 

Hair  and  hide 

59.26 

1.81 

6 30 

1 58 

0.058 

Offal  fat 

25.07 

69.44 

0.70 

0 32 

0.035 

Thoracic  fat 

Not  enoug 

h to  separa 

te. 

Total  animal 

62  43  1 

10.50 

3.35 

5.79 

1 008 

76 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  70. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 

(45-Months-Old.  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorus 

Animal  513,  Group  !. 

Lean  and  fat  flesh 

38.66 

49.37 

1.70 

0.51 

0.093 

Bone 

31.16 

20.30 

3.30 

25.14 

4.854 

Blood 

77.68 

3.43 

0.72 

0.028 

0.157 

Circulatory  system 

77.60 

4.50 

2.67 

0.91 

Respiratory  system 

73.91 

7.03 

2.74 

0.99 

0.155 

Nervous  system 

68.19 

15.50 

1.72 

1.60 

0.410 

Digestive  and  excretory  system 

72.23 

9.03 

2.44 

1.10 

0.194 

Hair  and  hide 

57.57 

10.07 

5.18 

0.94 

0.054 

Offal  fat 

5.68 

92.87 

0.22 

0.08 

0.011 

Thoracic  fat 

13.44 

83.71 

0.37 

0.19 

0.024 

Total  animal 

39.91 

42.85 

2.10 

3.20 

0.599 

Animal  502,  Group  II. 

Lean  and  fat  flesh 

61.63 

18.22 

2.91 

0.84 

0.156 

Bone 

33.00 

19.18 

3.35 

24.89 

4.408 

Blood  . . 

77.43 

3 49 

0 72 

0.023 

0.138 

Circulatory  system 

68.29 

14.51 

2.59 

0.68 

Respiratory  system 

76.17 

3.06 

2.83 

1.04 

0.166 

Nervous  system 

69.10 

14.60 

1.72 

1.79 

0.414 

Digestive  and  excretory  system 

75.18 

6.15 

2.59 

1.29 

0.174 

Hair  and  hide 

57.30 

3.10 

6.57 

0.98 

0.059 

Offal  fat 

10.65 

86.18 

0.47 

0.19 

0.023 

Thoracic  fat 

21.50 

73.48 

0.75 

0.30 

0.041 

Total  animal 

56.60 

16.97 

3.28 

5.09 

0.887 

Animal  509,  Group  III. 

Lean  and  fat  flesh 

63.17 

16.96 

2.89 

0.85 

0.160 

Bone 

33.59 

18.43 

3.42 

24.78 

4.466 

Blood  

78.05 

3.43 

0.69 

0 023 

Circulatory  system 

69.95 

12.26 

2.59 

0.92 

0.167 

Respiratory  system 

77.03 

2.88 

3.00 

1.01 

0.176 

Nervous  system 

66.87 

17.66 

1.68 

1.58 

0.379 

Digestive  and  excretory  system 

71.81 

10.58 

2.37 

1.01 

0.168 

Hair  and  hide 

58.97 

2.48 

6.27 

1.02 

0.046 

Offal  fat 

11.36 

85.89 

0.53 

0.18 

0.024 

Thoracic  fat 

19.81 

76.10 

0.72 

0.30 

0.043 

Total  animal 

57.72 

16.27 

3.23 

5.01 

0.887 

Table  71. — Composition  of  Certain  Parts  and  of  the  Total  Animal. 

(4- Year-Old  Cattle) 


Percent 

Water 

Percent 

Fat 

Percent 

Nitrogen 

Percent 

Ash 

Percent 

Phos- 

phorus 

Animal  501,  Group  1. 

Lean  and  fat  flesh 

36.25 

53.12 

1.46 

0.49 

0.087 

Bone 

32.09 

17.72 

3.36 

26.34 

4.808 

Blood 

77.98 

3.29 

0.86 

0.025 

Circulatory  system 

60.00 

24.55 

2.24 

0.74 

0.125 

Respiratory  system 

76.58 

3.35 

2.87 

1.04 

0.172 

Nervous  system 

70.40 

13.27 

1.67 

1.84 

0.392 

Digestive  and  excretory  system 

70.34 

12.06 

2.35 

0.95 

0.169 

Hair  and  hide 

51.43 

13.24 

5.49 

1.52 

0.049 

Offal  fat 

7.49 

91.06 

0.21 

0.10 

0.012 

Thoracic  fat 

18.83 

76.65 

0.61 

0.24 

0.026 

Total  animal 

38.75 

44.34 

2.00 

3.33 

0.587 

Animal  512,  Group  II. 

Lean  and  fat  flesh 

54.96 

28.29 

2.48 

0.77 

0.133 

Bone 

31.56 

20.31 

3.23 

25.62 

4.698 

Blood 

79.95 

3.07 

0.79 

0.023 

Circulatory  system 

54.88 

30.62 

2.14 

0.84 

0.117 

Respiratory  system 

75.49 

4.67 

2.80 

1.08 

0.164 

Nervous  system 

72.06 

11.12 

1.69 

1.87 

0.385 

Digestive  and  excretory  system 

72.04 

9.89 

2.36 

0.94 

0.164 

Hair  and  hide 

56.19 

3.61 

6.55 

1.16 

0.047 

Offal  fat 

11.21 

86.27 

0.37 

0.14 

0.019 

Thoracic  fat 

12.23 

85.52 

0.26 

0.14 

0.015 

Total  animal 

51.88 

24.09 

2.93 

5.10 

0.906 

Animal  500,  Group  III. 

Lean  and  fat  flesh 

63.11 

17.23 

2.96 

0.89 

0.156 

Bone 

33.05 

22.09 

3.19 

23.55 

4.208 

Blood 

79.04 

3.19 

0.79 

0.022 

Circulatory  system 

61.58 

22.27 

2.26 

0.80 

0.127 

Respiratory  system 

76.50 

2.70 

2.84 

1.15 

0.172 

Nervous  system 

62.77 

21.72 

1.67 

1.75 

0.371 

Digestive  and  excretory  system 

73.06 

9.29 

2.41 

1.00 

0.164 

Hair  and  hide 

59.33 

1.32 

6.28 

1.07 

0.044 

Offal  fat 

13.50 

83.56 

0.42 

0.18 

0.020 

Thoracic  fat 

17.04 

71.39 

0.49 

0.24 

0 024 

Total  animal 

57.25 

17.21 

3.20 

5.08 

0.884 

Studies  In  Animal  Nutrition — III 


77 


Table  72. — Composition  of  the  Total  Beef  Animal  on  Analytical,  Empty,  and 

Fat-Free  Bases. 


Group 

Age 

Basis 

% 

Water 

% 

Fat 

% 

Nitro- 

gen 

% 

Ash 

% 

Phos- 

phorus 

Embryo 

185  days 

Analytical 

84.801 

2.363 

1.673 

1.776 

0.283 

Embryo 

Embryo 

185  days 
232  days 

86.853 

1.713 

1.819 

0.290 

Analytical 

78.700 

2.573 

2.011 

3.180 

0.370 

Embryo 

Embryo 

232  days 
279  days 

Fat-free 

80.778 

2.064 

3.264 

0.380 

Analytical 

74.192 

3.384 

2.735 

4.062 

0.688 

Embryo 

Calves 

279  days 
at  birth 

Fat-free 

76.791 

2.831 

4.204 

0.712 

Analytical 

72.807 

3.648 

2.926 

4.523 

0.841 

Calves 

at  birth 

Live  weight 

73.583 

3.544 

2.843 

4.394 

0.817 

76.287 

2.947 

4.555 

0.847 

I 

3 months 

Analytical 

65.226 

9.712 

3.243 

4.875 

0.868 

I 

3 months 

Empty 

66.028 

9.488 

3.168 

4.763 

0.848 

I 

72.949 

3.500 

5.262 

0.937 

II 

3 months 

Analytical 

67.051 

7.348 

3.255 

4.971 

0.866 

II 

3 months 

Empty 

67.562 

7.234 

3.175 

4.894 

0.853 

II 

72.830 

3.422 

5.276 

0.919 

0.739 

III 

3 months 

Analytical 

71.108 

4.377 

3.247 

4.356 

III 

3 months 

Empty 

71.517 

4.315 

3.201 

4.295 

0.729 

III 

74.742 

3.345 

4.488 

0.761 

I 

5H  months 

Analytical 

56.771 

20.988 

2.753 

4.039 

0.731 

I 

5 Yi  months 

Empty 

57.213 

20.773 

2.725 

3.998 

0.723 

I 

5 Yi  months 
5lA  months 

Fat-free  empty 

72.214 

3.439 

3.130 

5.046 

0.913 

II 

Analytical 

63.966 

10.479 

4.996 

0.880 

II 

5M  months 

Empty 

64.388 

10.356 

3.093 

4.937 

0.870 

II 

5A  months 
5A  months 

Fat-free  empty 

71.827 

3.451 

5.508 

0.970 

III 

Analytical 

66.863 

6.884 

3.315 

4.947 

0.882 

m 

hxA  months 

Empty 

67.800 

6.689 

3.221 

4.807 

0.857 

hi 

hlA  months 
8H  months 

Fat-free  empty 

72.661 

3.452 

5.152 

0.919 

i 

Analytical 

61.009 

15.728 

2.952 

3.931 

0.704 

i 

8lA  months 

Empty 

61.233 

15.637 

2.935 

3.908 

0.700 

i 

8 A months 
8XA  months 

Fat-free  empty 

72.584 

3.479 

4.633 

0.830 

ii 

Analytical 

62.402 

13.936 

2.974 

4.389 

0.798 

ii 

8A  months 

Empty 

63.055 

13.695 

2.922 

4.312 

0.784 

ii 

8 M months 
8A  months 

Fat -free  empty 

73.060 

3.386 

4.997 

0.908 

m 

Analytical 

65.947 

8.590 

3.135 

5.010 

0.914 

hi 

8 XA  months 

Empty 

66.509 

8.437 

3.079 

4.921 

0.897 

hi 

8XA  months 
11  months 

Fat-free  empty 

72.637 

3.363 

5.374 

0.980 

i 

Analytical 

56.225 

21.078 

2.905 

3.862 

0.630 

i 

1 1 months 

Empty 

57.556 

20.437 

2.817 

3.744 

0.610 

i 

11  months 

Fat-free  empty 

72.340 

3.540 

4.706 

0.767 

i 

11  months 

Analytical 

53.228 

25.061 

2.785 

4.049 

0.749 

i 

11  months 

Empty 

54.424 

24.421 

2.714 

3.946 

0.730 

i 

11  months 

Fat-free  empty 

72.009 

3.591 

5.220 

0.966 

ii 

11  months 

Analytical 

61.651 

13.731 

3.076 

4.785 

0.799 

11 

11  months 

Empty 

63.006 

13.245 

2.967 

4.616 

0.771 

ii 

11  months 

Fat-free  empty 

72.626 

3.420 

5.320 

0.889 

ii 

11  months 

Analytical 

59.557 

16.361 

2.983 

4.969 

0.913 

ii 

11  months 

Empty 

60.371 

16.031 

2.923 

4.869 

0.895 

ii 

1 1 months 

Fat-free  empty 

71 . 897 

3.481 

5.799 

1.066 

in 

11  months 

Analytical 

62.925 

12.125 

3.068 

4.764 

0.846 

hi 

11  months 

Empty 

64.800 

11.512 

2.913 

4.532 

0.803 

in 

11  months 

Fat-free  empty 

73.231 

3.291 

5.111 

0.908 

78 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  73. — Composition  of  the  Total  Beef  Animal  on  Analytical,  Empty,  and 

Fat-Free  Bases. 


Group 

Age 

Basis 

% 

Water 

% 

Fat 

% 

Nitro- 

gen 

% 

Ash 

% 

Phos- 

phorus 

I 

18 

months 

Analytical 

51.695 

26.740 

2.748 

3.808 

0.680 

I 

18 

months 

Empty 

53.038 

25.997 

2.672 

3.702 

0.661 

I 

18 

71.671 

3.611 

5.003 

0.894 

III 

1,8J4  months 

Analytical 

62.641 

10.747 

3.258 

5.367 

0.950 

III 

l&A  months 

Empty 

65.411 

9.951 

3.017 

4.969 

0.879 

III 

72.639 

3.350 

5.518 

0.976 

I 

21 

months 

Analytical 

49.762 

29.420 

2.639 

4.015 

0.724 

I 

21 

months 

Empty 

50.933 

28.735 

2.577 

3.921 

0.708 

I 

21 

71.469 

3.616 

5.502 

0.993 

II 

26 

months 

Analytical 

60.368 

15.001 

3.095 

5.290 

0.897 

II 

26 

months 

Empty 

61.960 

14.398 

2.971 

5.077 

0.861 

II 

26 

72.382 

3.470 

5.931 

1.006 

III 

26 

months 

Analytical 

62.444 

11.871 

3.231 

5.088 

0.838 

III 

26 

months 

Empty 

64.115 

11.344 

3.087 

4.861 

0.801 

III 

26 

72.318 

3.482 

5.483 

0.903 

I 

34 

months 

Analytical 

43.684 

37.509 

2.286 

3.873 

0.614 

I 

34 

months 

Empty 

46.601 

35.566 

2.167 

3.672 

0.582 

l 

34 

72.324 

3.364 

5.699 

0.904 

II 

34 

months 

Analytical 

56. 102 

19.613 

3.031 

5.071 

0.792 

II 

34 

months 

Empty 

58.014 

18.759 

2.899 

4.850 

0.758 

II 

34 

months 

Fat-free  empty 

71.408 

3.568 

5.970 

0.933 

I 

39H  months 

Analytical 

38.628 

45.446 

2.019 

3.026 

0.507 

I 

3934  months 

Empty 

39.827 

44.558 

1.980 

2.967 

0.497 

1 

3Q14  months 

Fat-free  empty 

71.836 

3.571 

5.352 

0.897 

II 

40 

months 

Analytical 

55.333 

20.242 

3.039 

4.924 

0.877 

II 

40 

months 

Empty 

56.556 

19.688 

2.956 

4.789 

0.853 

II 

40 

months 

Fat-free  empty 

70.421 

3.680 

5.963 

1.062 

III 

4034  months 

Analytical 

62.430 

10.499 

3.353 

5.794 

1.008 

III 

4034  months 

Empty 

63.491 

10.202 

3.258 

5.630 

0.979 

III 

4034  months 

Fat-free  empty 

70. 705 

3.628 

6.270 

1.091 

I 

4434  months 

Analytical 

39.912 

42.850 

2.103 

3.201 

0.599 

I 

4434  months 

Empty 

41.396 

41.792 

2.051 

3.122 

0.585 

I 

4434  months 

Fat-free  empty 

71.117 

3.524 

5.363 

1.004 

11 

4434  months 

Analytical 

56.599 

16.972 

3.281 

5.086 

0.887 

II 

4434  months 

Empty 

57.625 

16.571 

3.203 

4.966 

0.866 

II 

44V4  months 

Fat-free  empty 

69.071 

3.840 

5.952 

1.038 

III 

45 

months 

Analytical 

57.715 

16.266 

3.234 

5.005 

0.887 

III 

45 

months 

Empty 

58.369 

16.015 

3.184 

4.928 

0.873 

III 

45 

months 

Fat-free  empty 

69.498 

3.791 

5.867 

1.040 

I 

47 

months 

Analytical 

38.752 

44.340 

1.999 

3.329 

0.587 

1 

47 

months 

Empty 

39.836 

43.556 

1.963 

3.270 

0.577 

I 

47 

months 

Fat-free  empty 

70.576 

3.478 

5.793 

1.022 

11 

48 

months 

Analytical 

51.879 

24.091 

2.927 

5.098 

0.906 

II 

48 

months 

Empty 

52.666 

23.697 

2.879 

5.014 

0.891 

II 

48 

months 

Fat-free  empty 

69.022 

3.773 

6.572 

1.167 

III 

48 

months 

Analytical 

57.254 

17.209 

3.200 

5.078 

0.884 

III 

48 

months 

Empty 

58.142 

16.852 

3.134 

4.972 

0.866 

III 

48 

months 

Fat-free  empty 

69.926 

3.769 

5.980 

1.041 

Studies  In  Animal  Nutrition — III 


79 


Table  74. — Composition  of  Beef  Flesh  on  Fat-Free  Basis. 


Group 

Age 

months 

Animal 

Sample 

% 

Water 

% 

Nitro- 

gen 

% 

Ash 

% 

Phos- 

phorus 

IU 

48 

500 

Round,  lean 

76.704 

3.236 

1.048 

0.198 

III 

48 

500 

Loin,  lean 

76.159 

3.374 

1.095 

0.201 

III 

48 

500 

Rib,  lean 

76.573 

3.645 

1.060 

0.194 

III 

48 

500 

Lean  and  fat  composite 

76.709 

3.511 

1.117 

0.196 

I 

47 

501 

Round,  lean 

77.117 

3.409 

1.056 

0.204 

I 

47 

501 

Loin,  lean 

76.228 

3.489 

1.037 

0.199 

1 

47 

501 

Rib,  lean 

75.900 

3.556 

1.019 

0.192 

I 

47 

501 

Lean  and  fat  composite 

76.537 

3.331 

1.182 

0.183 

II 

44  H 

502 

Round,  lean 

75.199 

3.446 

1.012 

0.203 

II 

44  H 

502 

Loin,  lean 

76.057 

3.475 

1.061 

0.217 

II 

44  H 

502 

Rib,  lean 

73.826 

3.413 

0.994 

0.182 

II 

44 

502 

Lean  and  fat  composite 

76.098 

3.616 

1.029 

0.195 

II 

11 

503 

Round  and  rump  lean 

76.776 

3.540 

1.076 

0.214 

II 

11 

503 

Loin,  lean 

77.548 

3.481 

1.074 

0.208 

II 

11 

503 

Rib,  lean 

77.618 

3.513 

1.033 

0.206 

I 

21 

504 

Round,  lean 

76.561 

3.533 

1.083 

0.214 

I 

21 

504 

Loin,  lean 

76.236 

3.476 

1.078 

0.206 

I 

21 

504 

Rib,  lean 

76.722 

3.565 

1.008 

0.202 

I 

11 

505 

Round  and  rump,  lean 

76.286 

3.674 

1.078 

0.221 

I 

11 

505 

Loin,  lean 

75.752 

3.595 

1.070 

0.218 

I 

11 

505 

Rib,  lean 

76.430 

3.668 

1.041 

0.218 

II 

34 

507 

Round,  lean 

77. 182 

3.428 

1.041 

0.204 

II 

34 

507 

Loin,  lean 

76.924 

3.040 

1.058 

0.201 

II 

34 

507 

Rib,  lean 

76.843 

3.417 

1.068 

0.198 

III 

45 

509 

Round,  lean 

76.862 

3.364 

1.049 

0.205 

m 

45 

509 

Loin,  lean 

76.527 

3.410 

1.045 

0.198 

hi 

45 

509 

Rib,  lean 

76.041 

3.283 

1.026 

0.091 

hi 

45 

509 

Lean  and  fat  composite 

76.359 

3.718 

1.039 

0.194 

ii 

48 

512 

Round,  lean 

76.770 

3.392 

1.073 

0.201 

ii 

48 

512 

Loin,  lean 

75.997 

3.458 

1.025 

0.191 

ii 

48 

512 

Rib,  lean 

76.566 

3.489 

1.053 

0.185 

ii 

48 

512 

Lean  and  fat  composite 

76.136 

3.496 

1.074 

0.188 

i 

44H 

513 

Round,  lean 

76.064 

3.387 

1.063 

0.202 

i 

44  H 

513 

Loin,  lean 

75.757 

3.542 

1.054 

0.208 

i 

44  M 

513 

Rib,  lean 

76.900 

3.551 

1.047 

0.196 

i 

44  H 

513 

Lean  and  fat  composite 

77.369 

3.345 

1.088 

0.194 

i 

34 

515 

Round,  lean 

76.102 

3.487 

1.039 

0.199 

i 

34 

515 

Loin,  lean 

76.599 

3.536 

1.172 

0.213 

i 

34 

515 

Rib,  lean 

76.851 

3.487 

1.066 

0.191 

ii 

26 

523 

Round,  lean 

78.748 

3.230 

1 067 

0.207 

ii 

26 

523 

Loin,  lean 

77.856 

3.379 

1.028 

0.192 

ii 

26 

523 

Rib,  lean 

77.486 

3.439 

1.022 

0.196 

80 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  75. — Composition  of  Beef  Flesh  on  Fat-Free  Basis. 


Group 

Age 

months 

Animal 

Sample 

% 

Water 

% 

Nitro- 

gen 

% 

Ash 

% 

Phos- 

phorus 

III 

40  M 

524 

Round,  lean 

79.010 

3.348 

1.072 

0.197 

III 

40H 

524 

Loin,  lean 

76.140 

3.468 

1.107 

0.203 

III 

40  H 

524 

Rib  lean  and  fat 

76.354 

3.596 

1.067 

0.198 

III 

40  H 

524 

Lean  and  fat  of  animal 

75.735 

3.509 

1.102 

0.201 

III 

26 

525 

Round,  lean 

78.931 

3.197 

1.090 

0.210 

III 

26 

525 

Loin,  lean 

77.316 

3.353 

1.086 

0.207 

III 

26 

525 

Rib,  lean 

77.199 

3.461 

1.070 

0.198 

II 

40 

526 

Round,  lean 

75.907 

3.745 

1.170 

0.217 

II 

40 

526 

Loin,  lean 

76.008 

3.391 

1.075 

0.201 

II 

40 

526 

Rib,  lean 

77.391 

3.419 

1.022 

0.183 

II 

40 

526 

Lean  and  fat  of  animal 

76.605 

3.595 

1.093 

0.186 

I 

39H 

527 

Round,  lean 

76.903 

3.477 

1.004 

0.203 

I 

39^ 

527 

Loin,  lean 

76.967 

3.518 

1.068 

0.203 

I 

39)4 

527 

Rib,  lean 

76.471 

3.542 

1.095 

0.196 

I 

39H 

527 

Lean  and  fat  of  animal 

76.535 

3.390 

1.073 

0.191 

I* 

38 

529 

Rib,  lean 

76.958 

3.474 

1.023 

0.192 

III 

18H 

531 

Round,  lean 

76.503 

3.333 

1.123 

0.212 

III 

18  H 

531 

Loin,  lean 

76.274 

3.460 

1.149 

0.213 

III 

18  H 

531 

Rib  lean  and  fat 

75.222 

3.611 

1.138 

0.206 

III 

18H 

531 

Lean  and  fat  of  carcass 

74.007 

3.686 

1.218 

0.227 

I 

18 

532 

Round,  lean 

75.867 

3.486 

1.124 

0.211 

I 

18 

532 

Loin,  lean 

75.664 

3.597 

1.111 

0.204 

I 

18 

532 

Rib,  lean 

76.008 

3.565 

1.106 

0.199 

I 

18 

532 

Lean  and  fat  of  carcass 

74.809 

3.747 

1.157 

0.209 

II 

11 

538 

Round,  lean 

78.078 

3.247 

1.123 

0.208 

II 

11 

538 

Loin,  lean 

77.494 

3.303 

1.118 

0.215 

II 

11 

538 

Rib,  lean 

77.170 

2.348 

1.107 

0.208 

II 

11 

538 

Lean  and  fat  of  carcass,  excl.  kidney  fat 

77.379 

3.379 

1.136 

0.205 

III 

11 

540 

Round,  lean 

77.444 

3.282 

1.107 

0.215 

III 

11 

540 

Loin,  lean 

76.759 

3.318 

1.122 

0.210 

III 

11 

540 

Rib  lean  and  fat 

76.867 

3.428 

1.122 

0.200 

III 

11 

540 

Lean  and  fat  of  carcass,  excl.  kidney  fat 

76.910 

3.304 

1.102 

0.191 

I 

11 

541 

Round,  lean 

76.490 

3.456 

1.127 

0.215 

I 

11 

541 

Loin,  lean 

76.187 

3.423 

1.112 

0.210 

I 

11 

541 

Rib,  lean 

76.654 

3.490 

1.090 

0.209 

I 

11 

541 

Lean  and  fat  of  carcass,  excl.  of  kidney  fat 

76.264 

3.686 

1.129 

0.212 

I 

8 H 

547 

Round,  lean 

77.207 

3.343 

1.119 

0.218 

I 

VA 

547 

Loin,  lean 

76. 177 

3.390 

1.097 

0.212 

I 

8A 

547 

Rib,  lean 

77.071 

3.360 

1.081 

0.199 

I 

8 H 

547 

Lean  and  fat  composite 

76.768 

3.348 

1.094 

0.197 

III 

5A 

548 

Round,  lean 

76.804 

3.420 

1.137 

0.218 

III 

5A 

548 

Loin,  lean 

76.878 

3.360 

1.151 

0.213 

III 

6H 

548 

Rib  lean  and  fat 

77.983 

3.221 

1.124 

0.200 

III 

5H 

548 

Lean  and  fat  composite 

77.576 

3.238 

1.095 

0.198 

III 

55 

549 

Rib,  lean 

77.060 

3.542 

1.164 

0.206 

♦Maintenance  one  year.  Then  full  feed. 


Studies  In  Animal  Nutrition — III 


81 


Table  76. — Composition  of  Beef  Flesh  on  Fat-Free  Basis. 


Group 

Age 

months 

Animal 

Sample 

% 

Water 

% 

Nitro- 

gen 

% 

Ash 

% 

Phos- 

phorus 

n 

8 H 

550 

Round,  lean 

77.563 

3.511 

1.092 

0.219 

II 

8 M 

550 

Loin,  lean 

77.442 

3.243 

1.109 

0.242 

II 

8 M 

550 

Rib  lean  and  fat 

77.651 

3.287 

1.120 

0.210 

II 

m 

550 

Lean  and  fat  composite 

77.704 

3.194 

1.056 

0.207 

n 

5y2 

552 

Round,  lean 

77.609 

3.320 

1.152 

0.211 

II 

5 'A 

552 

Loin,  lean 

77.334 

3.248 

1.142 

0.201 

II 

5 H 

552 

Rib,  lean  and  fat 

77.385 

3.455 

1.036 

0.192 

II 

5^ 

552 

Lean  and  fat  composite 

77.421 

3.237 

1.135 

0.187 

n 

3 

554 

Round,  lean 

77.629 

3.359 

1.299 

0.220 

ii 

3 

554 

Loin,  lean 

77.723 

3.322 

1.194 

0.215 

ii 

3 

554 

Rib,  lean  and  fat 

77.473 

3.378 

1.179 

0.205 

ii 

3 

554 

Lean  and  fat  composite 

77.986 

3.231 

1.257 

0.201 

hi 

3 

555 

Round,  lean 

79.013 

3.101 

1.210 

0.210 

HI 

3 

555 

Loin,  lean 

79.308 

3.160 

1.221 

0.212 

hi 

3 

555 

Rib,  lean  and  fat 

79.634 

3.182 

1.205 

0.194 

m 

3 

555 

Lean  and  fat  composite 

79.372 

3.150 

1.087 

0.193 

i 

3 

556 

Round,  lean 

77.660 

3.398 

1.292 

0.217 

i 

3 

556 

Loin,  lean 

77.099 

3.398 

1.236 

0.214 

i 

3 

556 

Rib,  lean  and  fat 

77.002 

3.432 

1.243 

0.198 

i 

3 

556 

Lean  and  fat  composite 

77.809 

3.333 

1.283 

0.196 

i 

5^ 

557 

Round,  lean 

77.392 

3.222 

1.060 

0.203 

i 

5 H 

557 

Loin,  lean 

77.196 

3.218 

1.191 

0.208 

i 

5 H 

557 

Rib,  lean 

77.069 

3.301 

1.088 

0.193 

i 

5 H 

557 

Lean  and  fat  composite 

77.549 

3.345 

1.292 

0.200 

m 

8H 

558 

Round,  lean 

78.020 

3.151 

1.061 

0.215 

m 

8H 

558 

Loin,  lean 

76.992 

3.269 

1.053 

0.216 

hi 

8H 

558 

Rib,  lean  and  fat 

77.379 

3.271 

1.026 

0.204 

hi 

sy 

558 

Lean  and  fat  composite 

77.830 

3.203 

1.006 

0.204 

Jersey 

6 days 

22A 

Flesh 

78.477 

3.743 

1.062 

0.204 

High  Plane 

Newborn 

562B 

Flesh 

79.108 

3.118 

1.110 

0.175 

High  Plane 

Newborn 

562C 

Flesh 

80.539 

2.724 

1.031 

0.180 

Medium  Plane 

Newborn 

565A 

Flesh 

80.318 

2.760 

0.951 

0.176 

Medium  Plane 

Newborn 

563A 

Flesh 

79.401 

2.921 

1.047 

0.163 

Medium  Plane 

Newborn 

564B 

Flesh 

79.925 

2.926 

0.989 

0.178 

Medium  Plane 

Newborn 

565B 

Flesh 

79.874 

2.909 

1.091 

0.187 

Medium  Plane 

Newborn 

564C 

Flesh 

79.302 

2.897 

1.071 

0.191 

Low  Plane 

Newborn 

568B 

Flesh 

82.931 

2.272 

0.896 

0.146 

Low  Plane 

Newborn 

567B 

Flesh 

80.332 

2.746 

1.119 

0.180 

Low  Plane 

Newborn 

566B 

Flesh 

82.039 

2.806 

1.049 

0.150 

Low  Plane 

Newborn 

568C 

Flesh 

79.293 

2.841 

1.057 

0.186 

High  and  1 
Medium  Plane / 

Newborn 

Average 

Flesh 

79.794 

2.902 

1.051 

0.179 

Low  Plane 

Newborn 

Average 

Flesh 

81.149 

2.666 

1.030 

0.166 

82 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  55 


Table  77. — Empty  Weight  at  Start. 


Animal 

Age 

atfirst 

feeding 

(days) 

Live  Weight 
at  first 
feeding 
(pounds) 

Live 

weight 

(kilograms) 

Empty 
weight 
(per  cent) 

Empty 

weight 

(kilograms) 

500 

32 

118.5 

53.750 

0.950 

51.1 

501 

10 

98.0 

44.452 

0.967 

43.0 

502 

15 

110.6 

50.167 

0.953 

47.8 

503 

23 

158.6 

71.939 

0.920 

66.2 

504 

21 

147.7 

66.995 

0.927 

62.1 

505 

21 

127.1 

57.651 

0.943 

54.4 

507 

19 

93.2 

42.275 

0.970 

41.0 

509 

15 

107.8 

48.897 

0.957 

46.8 

512 

28 

168.5 

76.430 

0.912 

69.7 

513 

14 

106.7 

48.398 

0.957 

46.3 

515 

19 

114.0 

51.709 

0.950 

49.1 

523 

23 

132.2 

59.965 

0.940 

56.4 

524 

19 

145.4 

65.952 

0.930 

61.3 

525 

36 

154.6 

70.125 

0.922 

64.7 

526 

41 

150.6 

68.311 

0.926 

63.3 

527 

27 

167.8 

76.112 

0.913 

69.5 

531 

73 

230.2 

104.416 

0.866 

90.4 

532 

54 

187.5 

85.048 

0.897 

76.3 

538 

25 

132.2 

59.965 

0.940 

56.4 

540 

32 

140.3 

63.639 

0.933 

59.4 

541 

20 

137.6 

62.414 

0.936 

58.4 

547 

12 

95.5 

43.318 

0.969 

42.0 

548 

20 

147.5 

66.905 

0.928 

62.1 

550 

21 

148.5 

67.358 

0.927 

62.4 

552 

18 

140.0 

63.503 

0.934 

59.3 

554 

18 

130.0 

58.967 

0.941 

55.5 

555 

21 

147.0 

66.678 

0.928 

61.9 

556 

19 

148.0 

67.131 

0.927 

62.2 

557 

23 

132.6 

60.146 

0.939 

56.5 

558 

13 

112.6 

51.074 

0.951 

48.6 

549 

18 

117.0 

53.070 

0.951 

50.5 

551 

21 

149.5 

67.812 

0.927 

62.9 

559 

19 

117.4 

53.251 

0.950 

50.6 

Table  78. — Amount  and  Composition  of  Gain  from  Start  to  Slaughter.  (Page  83) 


Weight 

Phos- 

phorus 

832.33 

521.86 
310.47 
665.91 

462.87 
203.04 

517.87 
518.72 

-0.85 

1,250.16 

471.78 

778.38 

864.21 

496.34 

367.87 
737.10 
521.02 
216.08 

1.199.78 
346.08 
853.70 

949.39 

523.54 
425.85 
807.60 
402.41 
405.19 

1.759.79 

488.22 
1,271.57 
2,002.23 

453.15 

1,549.08 

1,225.46 

470.94 

754.52 

2,116.15 

557.40 
1,558.75 
1,105.98 

497.18 

608.80 

3,035.37 

648.55 
2,386.82 

Percent 

Phos- 

phorus 

IlllsISllslllSlslilllglallsllsIlgSSgllsISsIsI 

00000000<J>000000000000000000000000000000000000 

Weight 

Ash 

4.673.7 

2.954.5 

1.719.2 
3,820.9 

2.614.1 

1.206.8 

3.052.6 

2.940.3 
112.3 

6.908.3 

2.661.2 

4.247.1 

4.905.1 
2,810.8 

2.094.3 

4.133.4 

2.949.8 

1.183.6 

6.700.4 

1.911.0 

4.789.4 

5.222.8 

2.970.2 

2.252.6 

4.428.8 

2.259.9 

2.168.9 
10,794.1 

2.762.3 

8.031.8 
10,824.9 

2.556.8 

8.268.1 

7.334.7 

2.656.4 
4,678.3 

11.512.8 

3.157.7 

8.355.1 

6.229.2 

2.815.6 

3.413.6 

16.994.9 
3,670.0 

13,324.49 

Percent 

Ash 

IgglSSlgllSlSSlIglillsIIlllsgsillssIlglsSlgal 

Weight 

Nitrogen 

3.108.8 

1.978.0 

1.130.8 

2.478.4 

1.742.7 
735.7 

2.274.9 

1.968.4 
306.5 

4.708.5 

1.779.8 

2.928.7 

3.073.2 

1.879.8 
1,193.4 

2.770.1 

1.974.8 
795.3 

5.031.8 

1.251.6 

3.780.2 

3.539.2 

1.984.3 

1.554.9 

2.771.2 

1.496.9 

1.274.3 
8,120.8 

1.845.4 

6.275.4 

7.445.3 

1.702.7 

5.742.6 

4.714.7 

1.776.6 

2.938.1 

6.910.6 

2.118.4 

4.792.2 

4.011.3 

1.883.0 

2.128.3 
12,264.6 

2.441.6 

9.823.0 

Percent 

Nitrogen 

SgSS§llSlgSllg|SSIIIillllllS§ISSllS|lllgSgglg 

COCOCOCOCOCOrOCOC*3<NCOC^COCOC^W(>DCOC^(>IC^CSlCOC^COCOeOC^COCaCO-OC^<NCOC^C<lCOC^CSICO(NC^COC^ 

Weight 

Fat 

9,310.6 

3.234.4 

6.076.2 

5.647.5 

2.664.0 

2.983.5 

3.067.3 
3,156.9 

-89.6 

35.895.9 

2.712.0 

33.183.9 
10,288.8 

2.965.0 

7.323.8 

5.751.6 
3,229.2 

2.522.4 

26.810.7 

1.680.0 

25.130.7 

16.585.7 

3.244.8 

13.340.9 

7.593.1 

2.138.4 

5.454.7 

58.918.3 
2,861.6 

56.056.7 

66.999.9 

2.556.8 
64,443.1 

21.048.5 

2.707.2 

18.341.3 
37,903.0 

3,508.6 

34.394.4 

15.855.4 

2.970.0 

12.885.4 

119.332.5 

4.578.0 

114.754.5 

Percent 

Fat 

lllSllsilcpIlIlll§ig!l§g|§§llSllll!lll3llsll 

Weight 

Water 

Percent 

Water 

l§ls§ISllsllI|gl§lslll§sllg|IISl§lllsggllllll 

Empty 

Weight 

98,133 

62,200 

35,933 

78,071 

55.500 
22,571 
71,078 
61,900 

9,178 

172,797 

56.500 

116.297 
99,349 

59.300 
40.049 
85,988 
62,100 
23,888 

171,488 

42,000 

129,448 

121,112 

62,400 

58,712 

89,999 

48,600 

41.399 

288.297 

58.400 
229,897 
274,357 

54.400 
219,957 
158,911 

56.400 
102,511 
236,429 

66,200 

170,229 

137,726 

59.400 
78,326 

459,025 

76.300 
382,725 

Condition 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

A„  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain  

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaugnter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

Animal 

556 

554 

555 

557 
552 
548 
547 
550 

558 
541 
505 
538 
503 
540 
532 

Age 

months 

£ £ £ ^ ^ £ 

e0e0C0*OM5U900  00  00-H»-l*-l>-l»-l00 

Group 

I 

m 

II 

II 
I 

I 

III 

II 

I 

III 

II 

I 

III 

II 
I 

(Page  84)  TABLE  78. — AMOUNT  AND  COMPOSITION  OF  GAIN  FROM  START  TO  SLAUGHTER — Continued. 


Weight 

Phos- 

phorus 

1,688.27 

777.44 

910.83 

3,366.90 

521.02 
2,845.88 

2.909.44 
470.94 

2,438.50 

2,126.98 

544.13 

1,582.85 

3,913.72 

406.55 

3,507.17 

3.174.44 

337.02 
2,837.42 
3,909.04 

584.05 

3,324.99 

3,651.97 

531.72 

3.120.25 
3,156.16 

513.69 

2,642.47 

4,507.90 

382.44 

4,125.46 

3,847.63 

395.31 
3,452.32 
3,418.30 

387.04 

3.031.26 
4,701.00 

354.32 
4,346.68 
4,399.23 

588.97 

3.810.26 
3,529.88 

422.47 

3,107.41 

Percent 

Phos- 

phorus 

SlIsllllllIglllKlsIIllIlgllllllsIgllslIIlllls 

OOOOOOOOOOOOOOOOOOOOOOOOOO'-'OOOOOOOOOOOOOOOOOO 

13 

9,540.5 

4.384.4 

5.156.1 
18,658.4 

2,949.8 

15.708.6 

17.150.6 

2.656.4 

14.494.2 

12.911.0 

3.079.7 

9.831.3 

24.673.2 

2.288.1 

22.385.1 

20.316.2 

1.857.3 
18,458.9 

23.320.7 

3.307.8 

20.012.9 
20,498.4 

3.013.1 

17.485.3 

18.141.7 

2.911.8 

15.229.9 

24.073.7 
2,134.4 

21.939.3 

22.068.9 

2.217.9 

19.851.0 

19.290.4 

2.162.2 

17.128.2 

26.645.1 

1.965.1 

24.680.0 

24.764.9 
3,338.6 

21.426.3 

20.279.1 

2.382.1 
17,897.0 

Percent 

Ash 

llliglsSSIIlBllllllggg|glBl2slIllIlllglsglslI 

Weight 

Nitrogen 

5.792.2 

2.892.8 

2.899.4 

12.264.1 

1.974.8 

10.289.3 

10.034.9 

1.776.6 

8.258.3 

8.199.5 

2.063.9 

6.135.6 

14.563.2 

1.512.3 

13.050.9 

12.142.4 

1.217.7 

10.924.7 

15.561.1 

2.214.4 

13.346.7 

12.649.9 

2.019.3 

10.630.6 

10.498.9 

1.949.3 

8.549.6 

15.819.3 
1,412.2 

14.407.1 

14.236.6 

1.467.5 

12.769.1 

12.464.4 

1.427.4 

11.037.0 

15.999.4 
1,290.0 

14.709.4 

14.217.6 

2.230.4 

11.987.2 

12.781.0 

1.577.9 

11.203.1 

Percent 

Nitrogen 

sllslISgllSlSlglllllllilSSlilllllSlIlllSlISsS 

Weight 

Fat 

19,105.6 

7,412.8 

11.692.8 

136.735.0 

3.229.2 

133.505.8 
48,636.5 

2.707.2 
45,929.3 

30.126.9 
3,429.1 

26.697.8 
238,975.5 

2,160.4 

236.815.1 

78.579.9 

1.640.0 

76.939.9 
350,228.8 

3.806.0 

346,422.8 

84.263.0 

3.291.6 

80.971.4 
32,874.7 

3.126.3 

29.748.4 
322,276.3 

1.944.6 
320,331.7 

73,645.3 

2.079.3 

71.566.0 
62,609.6 

2.012.4 

60.597.2 

354.940.0 
1,763.0 

353.177.0 
117,034.5 

3.833.5 

113.201.0 

68.726.2 

2.290.5 
66,436.1 

Percent 

Fat 

SlselllllsllllislIIllllllllalSslisllIsgllllls 

IS 

125,592 

60,026 

65,566 

242,366 

43,532 

198,834 

209,304 

40,044 

169,260 

170,280 

45,225 

125,055 

313,119 

35.254 

277,865 

243,018 

29,848 

213,170 

313,045 

47,956 

265,089 

242,058 

44,310 

197,748 

204,591 

43,094 

161,497 

319,219 

33,382 

285,837 

256,101 

34,416 

221,685 

228,490 

33,743 

194,747 

324,632 

31,089 

293,543 

260,103 

48,302 

211,801 

237,124 

36,444 

200,680 

Percent 

Water 

3iii§iiissiiiiiii§siisiisiiiiiiiiiiiiiiiiisii 

If 

192.005 

90.400 
101,605 
475,854 

62,100 

413,754 

337,803 

56.400 
281,403 
265,587 

64.700 
200,887 
671,917 

49,100 

622,817 

418.896 

41.000 

377.896 

786.005 
69,200 

716,805 

427,995 

63.300 
364,695 
322,234 

61.300 
260,934 
771,142 

46.300 
724,842 
444,424 

47.800 
396,624 
391,461 

46.800 
344,661 

814.914 

43.000 

771.914 
493,877 

69.700 
424,177 
407,833 

50.900 

356,933 

Condition 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain  

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain  

At  slaughter 

At  start 

Gam 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  start 

Gain 

At  slaughter 

At  3tart 

Gain 

At  slaughter 

At  start 

Gain 

Animal 

531 

504 

523 

525 
515 
507 
527 

526 

524 
513 
502 
509 
501 
512 
500 

Age 

months 

Group 

III 

II 

I 

III 

II 

I 

III 

II 

I 

II 

III 

II 

III 

Studies  In  Animal  Nutrition — III 


85 


Table  79. — Empty  Weight  at  Age  Previous  Animal  Was  Slaughtered. 


Animal 

Age 

Live  weight  at  end 
feeding 

Empty 
weight  at 
slaughter 

Per  cent 
empty 
weight 

Weight  at  age 
previous  animal  was 
slaughtered 

Yrs. — Mos. — Days 

lbs. 

kgs. 

kgs. 

Live 

kgs. 

Empty 

kg3. 

Group  1 


556 

0 

3 

0 

247.6 

112.31 

98.13 

87.38 

34.830 

557 

0 

5 

17 

443.2 

201.03 

172.80 

85.96 

109.41 

95.599 

547 

0 

8 

5 

450.2 

204.21 

171.45 

83.96 

142.93 

122.859 

541 

0 

10 

22 

724.0 

328.40 

288.30 

87.79 

250.38 

210.221 

505 

0 

10 

18 

690.6 

313.25 

274.36 

87.58 

255.37 

214.409 

532 

1 

5 

20 

1133.0 

513.92 

459.03 

89.32 

324.00 

284.439 

504 

1 

8 

26 

1170.0 

530.70 

475.85 

89.67 

488.74 

436.545 

515 

2 

9 

19 

1632.2 

740.35 

671.92 

90.76 

539.05 

483.363 

527 

3 

3 

15 

1869.0 

847.76 

786.01 

92.72 

777.45 

705.616 

513 

3 

8 

15 

1898.4 

861.10 

771.14 

89.55 

782.17 

725.229 

501 

3 

11 

0 

1964.8 

891.21 

814.91 

91.44 

873.71 

782.403 

Group  2 


554 

0 

3 

0 

196.0 

88.90 

78.07 

87.81 

34.830 

552 

0 

5 

7 

256.2 

116.21 

99.35 

85.49 

88.09 

77.349 

550 

0 

8 

14 

323.8 

146.87 

121.11 

82.46 

104.10 

88.994 

538 

0 

10 

26 

403.0 

182.80 

158.91 

86.93 

146.78 

121.036 

503 

0 

11 

11 

608.4 

275.96 

236.43 

85.67 

209.11 

172.428 

523 

2 

2 

6 

864.2 

391.99 

337.80 

86.18 

221.58 

192.619 

507 

2 

9 

16 

1014.4 

460.12 

418.90 

91.04 

440.12 

379.294 

526 

3 

4 

0 

1088.2 

493.60 

428.00 

86.71 

441.12 

401.592 

502 

3 

8 

19 

1138.6 

516.46 

444.42 

86.05 

491.69 

426.346 

512 

3 

11 

21 

1250.4 

567.17 

493.88 

87.08 

562.54 

484.067 

G^oup  3 


555 

0 

3 

0 

188.4 

85.46 

71.08 

83.17 

34.830 

548 

0 

5 

9 

222.0 

100.70 

85.99 

85.39 

79.02 

65.717 

558 

0 

8 

12 

237.8 

107.86 

90.00 

83.44 

94.12 

86.369 

540 

0 

11 

2 

341.8 

155.04 

137.73 

88.83 

135.90 

113.392 

531 

1 

6 

12 

479.6 

217.54 

192.01 

88.26 

153.99 

136.793 

525 

2 

2 

8 

694.6 

315.06 

265.59 

84.30 

240.49 

212.259 

524 

3 

4 

13 

806.2 

365.68 

322.23 

88.12 

299.60 

252.559 

509 

3 

8 

22 

1004.2 

455.50 

391.46 

85.94 

416.26 

366.808 

500 

3 

11 

26 

1061.8 

481.62 

407.83 

84.68 

438.30 

376.678 

(Page  86)  TABLE  80. — AMOUNT  AND  COMPOSITION  OF  GAIN  BETWEEN  EACH  SUCCEEDING  AGE. 


If 

.•8§S383ffS53g33!S8!Sl5SS8gS8S3S8SgS£§§S8§S8SS 

Per  cent. 
Phosphorus 

Is!SI!laill!lllIs§elalel§llS§IssSslllss 

OOOOOOOOOOOOOOOOOOOOt-hOOOOOOOO^hOOOOOOOOO 

b 

>NrfiWC0Tj<05Tj<05»0rHrl<NaiTH000)’^iC^05u:(NIN»0N(N»CNw:(NiHC0»0O<N00NNO 

Grams 

4,673. 

1,530. 

3,143. 

6,908. 

4,553. 

2,354. 

6,700. 

4,911. 

1.788. 

10,794. 

8,215. 

2,578. 

10,824. 

8,379. 

2,445. 

16,994. 

10,649. 

6,345. 

18,658. 

16,160. 

2,497. 

24,673. 

18,952. 

5,720. 

23,320. 

25,910. 

-2,589. 

24,073. 

21,517. 

2,556. 

26,645. 

24,426. 

2,218. 

26,567. 

25,910. 

656. 
24,073. 
24,512. 
— 439. 

Per  cent. 
Ash 

g21881SS§S8S3S8S31SS3SSSSE8SSlgS88ESS88 

^^^COT^COCOCOrOCOCOCOWW^COCOCOCOCOOCOCOCCWCOCOCOWiOMCOOCOWOCOCCO 

Weight 

Nitrogen 

.00NcD»0O05C0  0>05  00O00W05Tt(c0C0OiH»O«DNW05i-iN^W»0X^*HNNNOC0t0C0 

! si  Sill  §§  la  § IS  la  ill  S 11 11  ISIIsIIIIy  III.sII 

Per  cent. 
Nitrogen 

IllsssisSsllsIIisSsSIsiiSlsSIlSlISsslIsl 

Weight 

Fat 

.O^fMOi^iONiClNCCCOOOHGOiOOONOCO^tO^HX^^eOiOWOOlrHOi^iO  CO  <M  »-h 

Grams 

9,310 

1,234 

8,076 

35,895 

9,070 

26,825 

26,810 

25,521 

1,289 

58,918 

32.872 

26,046 

66,999 

33,527 

33,472 

119,332 

58,130 

61,201 

136.735 

113,488 

23,246 

238,975 

138,894 

100.081 

350,228 

250,959 

99,269 

322.276 
323,147 

—871 

354,940 

326,981 

27,958 

302,611 

250,959 

51,652 

322.276 
279,213 

43,063 

Per  cent. 
Fat 

13lR|S§Rl§sl§s|g3lslSlgllllSlllSllll»|g 

Weight 

Water 

Grams. 

64,795 

25,629 

39,166 

98,862 

63.122 

35.740 

104,983 

70,291 

34,692 

165,933 

128,725 

37,208 

149,316 

131,289 

18,027 

243,459 

163,712 

79,747 

242,366 

231,535 

10,831 

313,119 

246,191 

66,928 

313,045 

328,824 

—15,779 

319,219 

288,837 

30,382 

324,632 

323,884 

748 

341,755 

328,824 

12,931 

319,219 

315,330 

3,889 

Per  cent. 
Water 

lllal!lalSliSS!!lillSll!§Il!§S!!!ll!Ill 

Empty 

Weignt 

|gssKscsas|seg3S|sSi5|sggsg|sagsSgsg|sags 

Condition 

At  slaughter 

At  birth 

Gain 

At  slaughter 

At  3 mo 

Gain 

At  slaughter 

At  hVi  mo 

Gain 

At  slaughter 

At  8H  mo 

Gain 

At  slaughter 

At  8H  mo 

Gain 

At  slaughter 

At  11  mo  (541)... 

Gain 

At  slaughter 

At  18  mos 

Gain 

At  slaughter 

At  21  mos 

Gain 

At  slaughter 

At  34  mos 

Gain 

At  slaughter 

At  39}^  mos 

Gain 

At  slaughter 

At  21  mos 

Gain 

At  slaughter 

At  34  mos 

Gain 

At  slaugnter 

At  39^  mo 

Gain  

Animal 

556 

557 
547 
541 
505 
532 
504 
515 
527 
513 
501 
527 
513 

Age. 

months 

.^  = = 55Sii,n 
1 'iii1 1 1 1 i i 1 i 

Table  80. — Amount  and  Composition  op  Gain  Between  Each  Succeeding  Age — Continued.  (Page  87) 


If 

Per  cent. 
Phosphorus 

IsllllsSsRRKlssISIeillKlllsllSsISlgs 

OOOOOOOOOOOOOO'-IOOOOOOOOCSIOO’-IOOIMOOOOOIO 

Weignt 

Ash 

Grams 

3,820 

1,530 

2,290 

4,905 

3,785 

1,119 

5,222 

4,393 

8^9 

7,334 

5,219 

2,115 

11,512 

7,435 

4,077 

17,150 

8,891 

8,259 

20,316 

19,256 

1,059 

20,498 

19,477 

1,021 

22.068 

20,417 

1,651 

24,764 

24,038 

726 

19,821 

20,417 

—596 

24,764 

21,589 

3,175 

Per  cent. 
Ash 

IIISllsISsslllIssllsISlIlSsslllslIIS 

.^NNNCOTftNOONNOtOP5COO>C05^oo«005NN(OOOOOCONHO(X)(NONOi 

Weigh 

Nitroge 

Grams 

2,478 

990 

1,488 

3,073 

2,455 

617 

3,539 

2,752 

786 

4.714 
3,536 
1,178 
6,910 
5,038 
1,872 

10,034 

5.715 
4,319 

12,142 

11,268 

873 

12,649 

11,642 

1,007 

14.236 

12,602 

1,633 

14,217 

15,504 

—1,287 

12,755 

12,602 

152 

14,217 

13,892 

324 

a 

II 

SSIlSsSlSlgSlliSSllSsllllllslSIIlSIl 

IS 

T 

Weigh 

Fat 

Grams 

5,647 

1,234 

4,413 

10,288 

5,595 

4,693 

16,585 

9,216 

7,369 

21,048 

16,575 

4,472 

37,903 

23,614 

14,289 

48,636 

25,512 

23,124 

78,579 

54,610 

23,969 

84,263 

75,334 

8,928 

73,645 

83,939 

—10,293 

117,034 

80,214 

36,819 

90,218 

83,939 

6,279 

117,034 

98.265 

18,768 

Per  cent. 
Fat 

Weight 

Water 

Grams. 

52,746 

25,629 

27,117 

63,969 

52,259 

11,710 

76,367 

57,301 

19,066 

100.124 
76,319 
23,805 

142,735 

108,724 

34,011 

209,304 

121,362 

87,942 

243,018 

235.011 
8,007 

242,058 

232,980 

9,078 

256,101 

241.124 
14,977 

260,103 

278,944 

—18,841 

249.011 
241,124 

7,887 

260,103 

271,223 

—11,120 

er  cent. 
Water 

T T 

Empty 

Weight 

Grams. 

78.071 

34,830 

43,241 

99.349 

77.349 
22,000 

121,112 

88.994 

32,118 

158,911 

121,036 

37,875 

236,429 

172,428 

64,001 

337,803 

192.619 

145.184 

418,896 

379,294 

39,602 

427,995 

401,592 

26,403 

444,424 

426,346 

18,078 

493,877 

484,067 

9,810 

444,424 

426,346 

18,078 

493,877 

484,067 

9,810 

Condition 

At  slaughter 

At  birth 

Gain 

At  slaughter 

At  3 mo 

Gain 

At  slaughter 

At  5^  mo 

Gain 

At  slaughter 

At  8M  mo 

Gain  

At  slaughter 

At  8H  mo 

Gain 

At  slaughter 

At  11  mo. (538).. . 

Gain 

At  slaughter 

At  26  mo 

Gain 

At  slaughter 

At  34  mo 

Gain 

At  slaughter 

At  40  mo 

Gain  . 

At  slaughter 

At  44f%  mo 

Gain 

At  slaughter 

At  40  mos 

Gain 

At  slaughter 

At  44 Yi  mo 

Gain 

Animal 

554 

552 

550 

538 

503 

523 

507 

526 

502 

512 

502 

512 

months 

£ £ £ ^ 
-ill”"  'I 

(Page  88)  Table  80. — Amount  and  Composition  of  Gain  Between  Each  Succeedinq  Age — Continued. 


Weight 

Phosphorus 

Grams. 

517.87 

284.56 

233.31 

737.10 

479.08 

258.02 

807.60 

688.76 

118.84 

1.105.98 
1,017.13 

88.85 

1,688.27 

1,098.45 

589.82 

2.126.98 
1,865.76 

261.22 

3.156.16 
2,023.00 

1.133.16 
3,418.30 
3,591.05 
—172.75 
3,529.88 
3,288.40 

241.48 

2.738.99 
2,023.00 

715.99 

3,418.30 

3,117.87 

300.43 

Per  cent. 
Phosphorus 

SsISesISSlSIllIlllsIlSIglsSlIiSls 

O0000»-I00>-<00000»-H00000'-I00000000»-I00<-| 

Weight 

Ash 

,CD^N^i005acCU5NOCq»0H^OH0iN05  00^«05HN^O05H^oO 

CO  *0  OCOC0030<M  ^ ^ 2 § ^ § 22  ^ 52  SJ  ^ 2 £h  ^ 

Per  cent. 
Ash 

IllllSSssISIlllllSlISIlisllliSlgl 

Weight 

Nitrogen 

.Oi<Mr~^Heoir5cqf-»oeoeo®c<iooTt<i/5055C>a>iOTj<-«tieooo©'!t<50''*ioc&'^eo-H 

Grams 

2,274 

990 

1,284 

2.770 
2,103 

666 

2.771 
2,588 

182 

4,011 

3,491 

520 

5,792 

3,984 

1,807 

8,199 

6,403 

1.795 
10,498 

7.796 
2,702 

12,464 

11,950 

513 

12,781 

11,993 

787 

9,860 

7,796 

2,063 

12,464 

11,224 

1,240 

Per  cent. 
Nitrogen 

gI3SS5i&ggiSgSgggS8S83S13381583§g 

Weight 

Fat 

pnW^05C0NO'H05W^05iCOC0O05OONC0^tDW00«0OO^C000«0N^ 

Per  cent. 
Fat 

ss11s1S!1S§11b1II1II11111sI1SI11= 

'^CClO®-«f^OO«bg5^00«5©’^®^05«0©^«0«D®gcOCO©lCi-HOO©lC-H 

Weight 

Water 

Isssgs3ss-gs2sss§ssss3^||7s2ssss|a” 

Per  cent. 
Water 

BSl!ssi!g|!l31133iS2i!llSllISI!l! 

Empty 

Weight 

Grams. 

71,078 

34,830 

36,248 

85,988 

65,717 

20,271 

89,999 

80,369 

9,630 

137,726 

113,392 

24,334 

192,005 

136,793 

55,212 

265,587 

212,259 

53,328 

322,234 

252,559 

69,675 

391,461 

366,808 

24,653 

407,833 

376,678 

31,155 

322,234 

252,559 

69,675 

391,461 

366,808 

24,653 

Condition 

At  slaughter 

At  birth 

Gain 

At  slaughter 

At  3 mo 

Gain 

At  slaughter 

At  5)4  mo 

Gain 

At  slaughter 

At  8)4  mo 

Gain 

At  slaughter 

At  11  mo 

Gain 

At  slaughter 

At  18)4  mo 

Gain 

At  slaughter 

At  26  mo 

Gain 

At  slaughter 

At  40)4  mo 

Gain 

At  slaughter 

At  45  mo 

Gain 

At  slaughter 

At  26  mo 

Gain 

At  slaughter 

At  40)4  mo 

Gain 

Animal 

555 

548 

558 

540 

531 

525 

524 

509 

500 

524 

509 

months 

X X X ^ ^ 

eoiooo^ooookooooug 

1 1 J.  i 1 i T i T T | 

UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 
AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  56 


Observations  on  Winter  Injury 

I — Early  and  Late  Winter  Injury 

F.  C.  Bradford 

II — An  Aftermath  of  Winter  Injury 

H.  A.  Cardinell 


(Publication  Authorized  September  1,  1922.) 


COLUMBIA,  MISSOURI 
NOVEMBER,  1922 


UNIVERSITY  OF  MISSOURI 


COLLEGE  OF  AGRICULTURE 

Agricultural  Experiment  Station 


BOARD  OF  CONTROL 

THE  CURATORS  OF  THE  UNIVERSITY  OF  MISSOURI 

EXECUTIVE  BOARD  OF  THE  UNIVERSITY 
E.  LANSING  RAY  P.  E.  BURTON  H.  J.  BLANTON 

St.  Louis  Joplin  Paris 


ADVISORY  COUNCIL, 

THE  MISSOURI  STATE  BOARD  OF  AGRICULTURE 


OFFICERS  OF  THE  STATION 

J.  C.  JONES,  PH.  D.,  LL.  D.,  PRESIDENT  OF  THE  UNIVERSITY 
F.  B.  MUMFORD,  M.  S.,  DIRECTOR 


STATION  STAFF 

NOVEMBER,  1922 

AGRICULTURAL  CHEMISTRY 


RURAL  LIFE 


O.  R.  Johnson,  A.  M, 
S.  D.  GromEr,  A.  M. 
E.  L.  Morgan,  A.M. 


C.  R.  Moulton,  Ph.  D. 

L.  D.  Haigh,  Ph.  D. 

W.  S.  Ritchie,  Ph.  D. 

E.  E.  Vanatta,  M.  S. 

A.  R.  Hall,  B.  S.  in  Agr. 

E.  G.  Sieveking,  B.  S.  in  Agr. 

AGRICULTURAL  ENGINEERING 
J.  C.  Wooley,  B.  S. 

Mack  M.  Jones,  B.  S. 

ANIMAL  HUSBANDRY 

E.  A.  Trowbridge,  B.  S.  in  Agr. 

L.  A.  Weaver,  B.  S.  in  Agr. 

A.  G.  Hogan,  Ph.  D. 

F.  B.  Mumford,  M.  S. 

D.  W.  Chittenden,  B.  S.  in  Agr. 

A.  T.  Edinger,  B.  S.  in  Agr. 

H.  D.  Fox,  B.  S.  in  Agr. 

BOTANY 

W.  J.  Robbins,  Ph.  D. 

DAIRY  HUSBANDRY 
A.  C.  Ragsdale,  B.  S.  in  Agr. 

Wm.  H.  E.  Reid,  A.  M. 

Samuel  Brody,  M.  A. 

C.  W.  Turner,  B.  S.  in  Agr. 

D.  H.  Neison,  B.  S.  in  Agr. 

W.  P.  Hays 


ENTOMOLOGY 
Leonard  Haseman,  Ph.  D. 

K.  C.  Sullivan,  A.  M. 

O.  C.  McBride,  B.  S.  in  Agr. 

FIELD  CROPS 
W.  C.  Etheridge,  Ph.  D. 

C.  A.  Helm,  A.  M. 

L.  J.  Stadler,  Ph.  D. 

O W.  Letson,  B.  S.  in  Agr. 
Miss  Regina  Schulte* 


Ben  H.  Frame,  B.  S.  in  Aerr. 
Owen  Howells,  B.  S.  in  Agr. 


HORTICULTURE 
T.  J.  Talbert,  A.  M. 

H.  D.  Hooker,  Tr..  Ph.  D. 

J.  T.  Rosa,  Jr.,  Ph.  D. 

H.  G.  Swartwout.  B.  S.  in  Agr. 

J.  T.  Quinn,  B.  S.  in  Agr. 

POULTRY  HUSBANDRY 
H L.  Kempster.  B.  S. 

Earl  W.  Henderson,  B.S. 

SOILS 

M.  F.  Miller,  M.  S.  A. 

H.  H.  Krusekopf,  A.  M 
W.  A.  At  brecht.  Ph.  D. 

F.  L.  Duley,  A.M. 

Wm.  DeYoung,  B.  S.  in  Agr. 

H.  V.  Jordan,  B.  S.  in  Agr 
Richard  Bradfield,  Ph.  D. 

VETERINARY  SCIENCE 
J.  W.  Connaway,  D.  V.  S.,  M.  D. 

L.  S.  Backus,  D.  V.  M. 

O.  S.  Crisler,  D.  V.  M. 

A.  J.  Durant,  A.  M. 

H.  G.  Newman,  A.  M. 

OTHER  OFFICERS 

R.  B.  Price,  M.  S.,  Treasurer 
Leslie  Cowan,  B.  S.,  Secretary 

S.  B.  Shirkey,  A.  M.,  Asst,  to  Director 
A.  A.  Jeffrey,  A.  B.,  Agricultural  Editor 
J.  F.  Barham,  Photographer 

Miss  Jane  Frodsham,  Librarian. 

E.  E.  Brown,  Business  Manager. 


In  service  of  U.  S.  Department  of  Agriculture. 


Observations  on  Winter  Injury 

I. — Early  and  Late  Winter  Injury 

F.  C.  Bradford 

Destruction  of  flower  buds  without  attendant  damage  to  other 
parts  of  the  tree  is  the  most  common  manifestation  of  winter  injury 
in  the  peach  and  apparently  very  uncommon  in  the  apple.  Conversely, 
injury  to  other  tissues  without  attendant  damage  to  flower  buds  seems 
relatively  more  common  in  the  apple  than  in  the  peach.  In  short, 
blossom  buds  are  ordinarily  the  susceptible  point  in  the  peach  and 
the  resistant  part  of  the  apple  tree.  The  general  recognition  of  the 
connection  between  responsiveness  of  peach  flower  buds  to  high  temper- 
atures in  winter  and  the  too  frequent  consequent  damage  from  ordi- 
nary winter  cold  has  fostered  a rather  widespread  assumption  that 
injury  to  flower  buds  of  any  kind  must  be  attributed  to  this  combin- 
ation of  weather  conditions.  For  this  reason  an  occurrence  of  injury 
confined  to  blossoms  in  the  apple  while  peaches  in  the  same  orchards 
were  wholly  undamaged,  constituting  a case  of  injury  supposedly  due 
to  premature  starting  from  dormacy  in  a fruit  supposedly  least  sub- 
ject to  this  weakness,  merits  investigation  and  record. 

MANIFESTATIONS 

Late  in  December,  1921,  it  was  noted  that  a small  percentage 
of  fruit  buds  on  Jonathan  trees  in  the  University  orchard  at  Columbia 
had  been  injured,  the  floral  parts  being  clearly  discolored.  In  the 
spring  of  1922  these  trees  blossomed  very  heavily.  Shortly  after  the 
blossoms  had  fallen,  the  unusual  persistence  of  the  bud  scales  and 
the  peculiar  behavior  of  the  axillary  buds  attracted  attention.  In 
many  cases  the  terminal  buds  were  dead  and  growth  was  proceeding 
from  axillary  buds.  Most  of  the  dead  buds  abscissed,  leaving  a smooth, 
flat  surface,  as  shown  in  Fig.  1.  In  other  cases  the  terminal  growth 
was  very  feeble  and  fruit  had  set  from  axial  blossom  clusters.  An  in- 
stance of  this  is  shown  in  Fig.  2.  Since  in  Jonathan  the  formation  of 
axillary  flower  buds  without  accompanying  flower  buds  on  terminals 
has  not  been  observed,  the  association  of  axillary  blossoms  with  leafy 
terminals  invited  attention.  Furthermore,  the  setting  of  fruit  from 
axillary  buds  in  such  numbers  as  appeared  this  spring  was  unusual ; 
ordinarily  the  terminal  buds  set  fruit  and  the  axillary  buds  do  not. 


4 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


Many  spurs  which  had  not  blossomed  this  year  bore  fewer  leaves 
than  is  usual  in  ncn-blossoming  spurs.  Figs.  3 and  4 show  views  from 
two  angles  of  a spur  of  this  type  found  on  a Ben  Davis  tree.  Careful 
examination  of  either  reveals  a sharply  marked  ring  just  above  the 
lowest  leaves.  An  enlarged  view  of  a similar  spur  (Fig.  5)  shows 
the  significance  of  this  appearance  perhaps  more  clearly.  Here  the 
ring,  just  below  the  leaf  insertions,  is  plainly  visible.  This  ring,  a 
single  continuous  line,  is  quite  different  from  the  composite  belt  of 
scale  scars  marking  the  transition  from  one  year’s  growth  to  the  next. 
It  occurs  only  on  fruiting  wood  at  the  point  where  the  vegetative  axis 
leaves  the  purse. 

At  the  right  of  the  spur,  just  below  the  ring,  is  a black  protuber- 
ance; this  is  the  dried  remnant  of  a flower  cluster  which  was  killed 
in  the  bud.  The  purse,  or  cluster  base,  the  swelling  which  bears  the 
blossoms  and  fruit,  so  prominent  in  the  majority  of  bearing  spurs 
(cf.  Fig.  15),  is  here  reduced  to  extremely  small  dimensions.  The 
vegetative  axis,  relatively  small  in  many  blossoming  or  bearing  spurs, 
in  these  cases  constitute  nearly  all  the  current  season’s  growth.  In 
some  the  purse  is  reduced  to  even  smaller  size  and  can  scarcely  be  dis- 
tinguished (Fig.  8).  In  others  it  is  fairly  conspicuous. 

Fig.  6 is  a photomicrograph  of  a longitudional  section  of  the  bud 
shown  in  Fig.  5.  Though  this  section  does  not  pass  through  the  exact 
center,  the  dead  blossoms  within  the  cluster  are  discernible.  The 
smallness  of  the  pith  cylinder  extending  to  the  blossoms  is  rather  bet- 
ter evidence  than  the  possibly  shrivelled  blossoms  could  be,  of  the 
size  of  the  bud  at  the  time  of  the  killing.  Fig.  7,  representing  a Jan- 
uary stage  in  an  uninjured  bud  of  another  variety,  is  used  here  to  illus- 
trate the  extent  of  the  injury  to  the  Jonathan  bud.  That  part  of  the 
bud  represented  by  the  portion  above  the  line  k k’  was  killed.  Growth 
continued  from  the  vegetative  point  just  below  this  (v),  resulting  in 
the  growth  shown  to  the  left  of  and  above  the  injured  portion. 

A section  of  another  bud  is  shown  in  Fig.  9.  In  this  case  the  dead 
tissue  had  abscissed  and  the  pith  cylinder  extending  to  the  point  of 
abscission  (a)  is  the  best  evidence  of  the  former  presence  of  blossoms 
at  this  point.  Injury  to  the  pith  is  shown  at  k and  the  growth  from 
the  vegetative  point  was  less  vigorous.  More  severe  injury  is  shown 
in  Fig.  10,  also  some  regenerative  tissue.  Complete  killing  is  shown 
in  Fig.  11  and,  in  Fig.  12,  complete  killing  followed  by  abscission. 

Dissection  of  these  twigs  and  spurs  showed,  in  many  cases,  though 
not  invariably,  some  discoloration  of  the  pith.  Generally  when  the 
bud  had  sloughed  off,  as  shown  in  Fig.  1,  little  injury  was  visible  in 


Observations  on  Winter  Injury 


Plate  I. — Fig.  1.  Jonathan,  showing  abscission  of  winter-killed  terminal.  Fig. 
2.  Jonathan,  showing  setting  of  fruit  from  axial  blossom  clusters.  Figs.  3 and  4.  Ben 
Davis  spur,  showing  winter  killing  of  blossoms.  Fig.  5.  Jonathan  spur,  showing 
dead  blossoms.  Fig.  6 Photomicrograph  of  section  of  spur  shown  in  Fig.  5. 


6 


Missouri  Agr.  Exp.  Sta.  Research  Bullet 


56 


Plate  IT. — Fig.  7.  Uninjured  bud.  In  injured  buds,  tissues  above  k-k'  killed 
and  growth  proceeds  from  v.  Fig.  8.  Jonathan  spur,  showing  dead  blossoms  at  k and 
extreme  reduction  of  purse.  Fig.  9 Jonathan  spur,  showing  abscission  of  dead 

blossoms  at  a and  injury  to  pith  at  k.  Fig  10.  Similar  injury  involving  greater  area. 
Fig.  11.  Whole  bud  killed.  Fig.  12,  bud  killed  and  abscissed. 


Observations  on  Winter  Injury 


7 


Plate  III. — Fig.  13.  Kicffer  pear,  showing  replacement  of  killed  spurs 
from  supernumerary  buds.  Fig.  14.  Kieffer  pear  spur,  injured  in  spring  of 
1921,  bearing  in  1922.  Injury  shown  in  blackened  wood.  Fig.  15.  Ben 
Davis  spur,  July,  1921.  Bearing  from  normal  and  from  second  bloom. 


%ai 

Fig.  13 


8 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


Plate  IV. — Series  of  sections  of  Kieffer  pear.  Fig.  16.  Pith  injury  near 
tip  of  1921  wood.  Fig.  17.  Injury  to  pith  and  wood  in  1920  wood.  Fig.  18. 
Section  from  lower  level  in  1920  wood;  more  wood  and  less  pith  injury. 
Fig.  19.  Still  lower  on  1920  wood;  injury  confined  to  wood. 


Observations  on  Winter  Injury 


9 


the  remaining  tissue.  Some  of  the  spurs  in  which  the  blossoms  had 
been  killed  showed  no  further  injury.  In  the  majority  of  these  cases, 
however,  there  were  rather  poorly  defined  small  areas  in  which  the 
pith  was  in  some  cases  browm,  in  some  cases  orange.  Other  spurs, 
however,  which  were  bearing  normally  showed  somewhat  the  same 
injury.  This  has  been  observed  in  Oregon  and  in  Missouri  in  other 
seasons. 

Parenthetically  it  may  be  recorded  that  spurs  somewhat  similar 
in  external  appearance  at  this  time  to  those  just  described,  but  with 
the  purse  better  developed,  were  showm  by  dissection  to  be  infected 
with  fire  blight.  Here  the  discoloration  was  black  and  primarily  in  the 
vessels  rather  than  the  pith.  Its  course  from  the  blossom  attachments 
was  easily  traced.  Leaves  on  infected  spurs  at  this  stage  are  still 
green  and  practically  the  only  external  evidence  of  infection  is  the 
rather  flaccid  condition  of  the  purse. 

OCCURRENCE 

The  distribution  of' this  winter  injury  was  rather  general  among 
the  Jonathan  trees  in  two  parts  of  the  University  orchard.  Other 
varieties  examined,  as  Ben  Davis,  Ingram  and  Wealthy,  showed  it,  but 
very  rarely.  Others,  as  Oldenburg,  York  and  Gano,  apparently  had 
none.  It  was  very  common  in  Malus  prunifolia.  In  Jonathan  it  affected 
certainly  10  per  cent  of  the  buds.  It  was  found  in  young  Jonathans 
in  the  cultivated  orchard  at  Turner  though  less  abundantly  than  in  the 
sod  orchard  in  Columbia.  Possibly  the  trees  in  the  better  drained 
positions  suffered  less,  but  the  differences  were  not  marked.  In  a row 
on  the  northern  boundary  of  the  orchard  injured  buds  were  distinctly 
more  plentiful  on  the  north  side  of  the  trees.  This  might  have  been 
taken  as  evidence  of  a cold  wind  as  a factor  in  the  injury,  were  it  not 
that  elsewhere  in  the  orchard  injury  was  distinctly  localized  on  other 
sides.  Almost  invariably  it  was  most  abundant  on  that  side  which 
faced  an  open  space,  regardless  of  orientation. 

Much  more  consistent  was  the  preponderance  of  injury  in  buds 
on  terminal  shoots  over  that  on  spurs.  Though  spurs  greatly  outnum- 
ber terminals,  probably  three-fourths  of  the  cases  of  injury  appeared 
on  terminal  shoots.  The  importance  of  this  observation  will  appear 
later. 

In  the  peach  no  sign  of  injury  could  be  found,  though  many  axil- 
lary leaf  buds  failed  to  open  in  the  spring.  Poorly  developed  axillary 
buds  on  vigorous  shoots  of  many  perennials,  whether  they  are  those 


10 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


formed  in  the  first  flush  of  spring  growth  or  those  laid  down  just  be- 
fore growth  stops  in  late  autumn,  are  inclined  to  remain  dormant. 
Consequently  this  failure  to  grow  is  no  indication  of  winter  injury, 
particularly  in  view  of  the  absence  of  any  evidence  of  injured  tissue. 
In  the  peach  the  pith  was  brown  in  spots,  but  the  cells  were  not  col- 
lapsed. In  this  fruit  the  pith  is  short-lived  and  discoloration  often 
occurs  in  new  growth  before  midsummer.  Fruit  buds  were,  practically 
without  exception,  uninjured.  Consequently  it  may  be  said  that  the 
peach  came  through  the  winter  without  injury. 


Table  1. — Precipitation  and  Temperature  at  Columbia,  September  to  Decem- 
ber, 1921,  Compared  to  That  oe  the  Same  Months,  1911  to  1920  Inclusive. 

Precipitation  (inches)  Aug  Sept.  Oct.  Nov.  Dec. 


1921  5.83 

Average,  1911-1920  4.90 

Maximum  7.83 

Minimum  0.77 


Daily  Mean  Temperature  (°F.) 

1921  

Average,  1911-1920  

Maximum  (monthly)  

Minimum  (monthly)  

Absolute  Minimum  Temperature  (°F.) 

1921  

1911-1920  


10.04 

2.33 

1.34 

1.41 

6.06 

4.90 

1.86 

1.63 

9.69 

7.68 

3.24 

2.94 

2.38 

0.72 

0.10 

0.44 

71.9 

57.4 

43.8 

35.5 

68.6 

57.1 

45.1 

32.4 

73.0 

60.1 

48.9 

39.8 

61.4 

48.5 

38.0 

25.4 

44 

32 

17 

7 

32 

20 

6 

—9 

CAUSE 

The  records  of  the  United  States  Weather  Bureau  Station  at  Co- 
lumbia, made  available  by  Dr.  George  Reeder,  section  director  for 
Missouri,  are  summarized  in  Table  1,  to  December  31,  covering  the 
period  during  which  the  injury  occurred.  They  show  the  mean  tem- 
peratures for  September  and  December  to  have  been  rather  well  above 
the  averages  for  the  previous  ten  years  and  that  for  November  some- 
what lower,  but  in  no  case  did  they  reach  the  extremes  recorded  in  the 
previous  ten  years.  The  absolute  minima  for  the  several  months  have 
in  each  case  been  exceeded  in  other  recent  years.  Unusual  cold  is 
evidently  not  the  primary  factor  in  this  injury. 

The  precipitation  record,  however,  shows  an  unusual  rainfall  in 
September,  higher  than  any  in  the  previous  ten  years;  indeed  it  was 
exceeded  but  twice  in  35  years.  This  rainfall,  coupled  with  the  con- 
siderable increase  in  mean  temperature  for  the  same  month,  undoubt- 


Observations  on  Winter  Injury 


11 


edly  tended  to  delay  maturity.  Raspberries  made  considerable  new 
growth  and  fall  blossoming  of  cherries  was  rather  widespread.  Some 
Japanese  plums  in  the  University  orchard  were  partly  in  blossom  in 
October.  Though  the  subsequent  winter  was  not  at  all  remarkable  for 
cold  weather,  in  either  duration  or  intensity,  it  resulted  in  practically 
complete  destruction  of  red  and  purple  raspberry  canes  and  serious 
damage  to  black  raspberries. 

Injury  confined  to  flower  buds  during  early  winter  has  been  re- 
corded but  rarely.  Maynard4*  and  Bartlett1  described  cases  of  Decem- 
ber killing  of  peach  buds  in  Massachusetts.  In  the  case  cited  by 
Maynard  the  cold  was  not  intense  (10°F.),  but  it  followed  warm 
weather  during  which  the  blossoms  were  observed  to  develop.  Chand- 
ler3 found  mild  injury  to  peach  blossom  buds  in  New  York  during 
the  winter  of  1914-1915.  The  previous  August  had  been  character- 
ized by  heavy  precipitation.  The  coldest  temperature  of  the  winter, 
— 9°F.,  occurred  in  December.  The  injury  was  in  the  pith  of  the  bud 
and  of  the  twig  and  apparently  had  no  effect  beyond  retarding  blos- 
soming. In  these  cases  it  is  not  altogether  clear  whether  the  buds  had 
started  to  develop  or  had  failed  to  mature. 

The  only  careful  report  of  winter  injury  to  blossom  buds  in  apple 
unaccompanied  by  further  injury  is  that  of  a case  observed  by  Whip- 
ple5 in  Montana.  This  form  is  apparently  identical  in  its  manifesta- 
tions with  that  just  described  for  Columbia.  Though  this  form  of 
injury  seems  rare,  it  is  quite  possible,  as  Whipple  points  out,  that, 
since  the  injury  escapes  casual  observation,  it  may  occur  from  time 
to  time  and  pass  unnoticed.  Whipple  suggested  that  the  damage  in 
Montana  might  have  been  due  to  thawing  in  high  winds  or  to  freezing 
after  warm  weather  in  January  or  February. 

All  the  evidence  in  the  occurrence  at  Columbia,  however,  con- 
nects immaturity  with  the  injury.  It  followed  weather  conditions  in- 
viting and  in  many  cases  leading  directly  to  immaturity  injuries  in  other 
fruits.  Peaches,  far  more  susceptible  to  injury  from  premature  de- 
velopment, were  entirely  uninjured.  The  greater  injury  on  open 
sides  may  have  been  due  to  prolonged  growth  as  much  as  to  greater 
exposure.  In  the  varieties  affected,  damage  was  greatest  in  the  ter- 
minals on  shoots;  these  are  late  in  differentiating  flower  buds,  late  in 
maturing  and  late  in  starting  from  dormancy.  Some  late  blossoming 
varieties  were  affected  while  some  early  blossoming  varieties  escaped. 
Finally,  the  injury  had  occurred  before  the  last  of  December. 

•This  and  subsequent  superscript  numerals  refer  to  literature  cited  in  the  Bibliography. 


12 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


IMPLICATIONS 

Attention  of  fruit  growers  in  Missouri  has  been  focused  on  pre- 
mature starting  of  buds  during  winter  rather  than  on  immaturity.  In 
this  case  peach  trees  in  the  same  orchard  showed  no  injury,  either  in 
wood  or  in  fruit  bud.  In  addition  to  weather,  other  conditions  were 
particularly  favorable  to  injury  from  immaturity  in  the  peach  this  win- 
ter, for,  following  the  Easter  freeze  in  1921  the  trees  had  been  cut 
back  to  wood  2 or  3 inches  in  diameter  and  had  made  growths  of  three 
to  6 or  7 feet,  with  secondary  and  even  tertiary  branches.  Consequently 
their  immunity  from  injury  while  Jonathan  apples  were  afflicted  indi- 
cates that  in  the  peach  at  Columbia  immaturity  is  less  important  than 
premature  starting.  On  the  other  hand,  so  far  as  concerns  the  apple, 
evidence  is  accumulating  in  a chain,  of  which  the  case  here  recorded 
is  but  one  link,  that  even  to  Central  Missouri,  as  is  the  case  farther 
north,  immaturity  is  of  no  mean  importance.  Crown  rot  in  Grimes 
and  crotch  injury  in  Stayman  are  presumably  due  to  injury  consequent 
upon  immaturity.  Cardinell  has  found  serious  cases  of  winter  injury — 
with  heart  rot  as  the  consequence — in  young  apples,  evidently  tracing 
to  immaturity  and  cold  in  the  fall  of  1917.  Injuries  due  to  immaturity 
are  not  patent.  The  type  described  here  might  well  escape  observation 
and  the  crop  failure  be  attributed  to  lack  of  fruit  bud  formation.  In- 
juries to  other  tissues  are  often  unnoted  until  brought  to  attenion  by 
the  wood-destroying  fungi  which  find  entrance  through  such  lesions, 
or  until  the  bark  comes  away,  long  after  the  weather  conditions  re- 
sponsible for  the  injury  have  passed  from  recollection.  In  many  cases 
such  injuries  are  attributed  to  fungi  and  referred  to  under  the  general 
term  “canker’. 

If  the  conventional  notions  of  the  effects  of  cultivation  be  accepted, 
peaches  and  apples  may  be  bad  neighbors  in  Central  Missouri  orchards, 
for  the  cultural  practices  which,  by  inducing  late  growth,  tend  to  make 
peaches  resist  stimulation  from  warm  weather  in  January  and  Feb- 
ruary tend  to  make  some  apples  more  subject  to  injury  in  November 
and  December.  When  peaches  are  used  as  fillers  in  apple  orchards, 
cultural  practices  designed  to  protect  either  fruit  against  winter  in- 
jury are  likely  to  make  the  other  more  susceptible. 

However,  it  should  be  recognized  that  the  comparison  made  here 
is  between  peaches  in  cultivated  soil  and  apples  in  sod.  The  generally 
accepted  views  of  the  effects  of  these  two  systems  of  management  are, 
for  the  most  part,  founded  on  investigations  in  sections  with  a shorter 
and  uniform  growing  season.  For  many  crops  there  are  in  Missouri 


Observations  on  Winter  Injury 


13 


two  rather  distinct  growing  seasons,  separated  by  a season  of  dry,  hot 
weather.  In  some  cases,  as  with  potatoes  and  cabbage,  this  may  be 
due  chiefly  to  the  excessively  high  temperature  of  midsummer,  but 
with  others  dry  weather  has  its  undoubted  influence.  The  raspberry, 
companion  of  the  potato  and  the  cabbage  in  ability  to  grow  where  the 
summers  are  too  cool  to  ripen  grain,  shows  the  same  reaction  to  the 
growing  season  of  Central  Missouri.  Immaturity  injury  in  this  fruit 
occurs  every  year  in  varying  degree  at  Columbia  before  intense  cold 
sets  in ; it  may  be  produced  by  temperatures  certainly  no  lower  than 
12 °F.  and  possibly  higher.  In  late  August  the  canes  are  often  more 
nearly  mature  than  they  are  in  October,  following  the  moderate  tem- 
perature and  greater  rainfall  of  September.  Those  which  grow 
through  August  seem  to  mature  better  than  those  which  stop  growing 
at  this  time.  Card2  found  in  New  York  that  under  some  circumstances 
the  first  shoots  to  start  in  the  spring  may  be  more  tender  in  the  follow- 
ing winter  than  those  starting  somewhat  later.  Prolonged  tillage 
through  the  dry  season  may  have  the  actual  effect  of  inducing  final 
maturity  by  so  prolonging  the  first  flush  of  growth  that  the  second 
growth  does  not  start.  In  short,  for  this  fruit  the  growing  season  here 
is  apparently  too  long;  the  canes  mature  and  then  resume  growth. 

On  the  other  hand,  the  peach,  adjusted  to  warmer  summers,  suf- 
fers little  or  no  check  from  heat  during  its  growing  season.  The  long 
period  of  warm  weather  enables  this  tree  to  mature  properly  despite 
high  cultivation.  In  fact,  prolonged  cultivation  makes  it  more  hardy. 

The  apple,  with  growing  season  temperature  requirements  higher 
than  those  of  the  raspberry  and  lower  than  those  of  the  peach,  un- 
doubtedly suffers  more  or  less  here  from  immaturity.  The  evidence 
at  hand,  however,  does  not  warrant  any  conjecture  as  to  the  effects  of 
tillage  on  maturity  in  this  fruit.  The  smaller  amount  of  injury  in  the 
cultivated  trees  at  Turner  than  in  the  sod  orchard  at  Columbia,  eight 
miles  away,  is  interesting,  possibly  suggestive,  but  certainly  not  indi- 
cative. The  greater  prevalence  of  injury  on  terminals  than  on  spurs 
at  Columbia  points  in  the  opposite  direction.  Consequently  at  present 
it  cannot  be  stated  definitely  whether  this  immaturity  injury  is  due  to 
prolonged  growth  or  to  renewed  growth. 

One  thing  becomes  increasingly  evident.  Hardiness  in  Central 
Missouri  is  more  complicated  than  it  is  farther  north  or  farther  south. 
In  some  regions  it  is  largely  a matter  of  maturity,  in  others  a matter 
of  continuing  dormancy.  Here  it  is  in  some  fruits  the  first,  in  other 
fruits  the  second.  This  is  the  first  complication.  The  second  compli- 
cation comes  from  the  fact  that  immaturity  alone,  a rather  simple  mat- 


14 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


ter  farther  north,  is  here  possibly  induced  by  the  very  practices  that 
obviate  it  elsewhere.  Finally,  since  this  section  shares  northern  and 
southern  winter  weather,  extreme  measures  for  guarding  against  one 
type  of  injury  may  be  efficacious  in  one  winter  and  injurious  in  the 
next.  Solution  of  the  problems  raised  will  depend  on  recognition  of 
the  types  of  injury  to  which  each  fruit  is  subject,  determination  of  the 
probability  of  the  occurrence  of  weather  conditions  leading  to  each 
type  and  formulation  for  each  fruit  of  cultural  practices  which,  over  a 
period  of  years,  will  reduce  the  injuries  most  likely  to  occur. 

VARIABILITY  OF  HARDINESS 

After  the  Easter  freeze  of  1921  the  Kieffer  pears  showed  more 
damage  than  any  other  trees  in  the  University  orchard.  Many  spurs 
were  killed,  many  branches  were  killed  back  well  into  1920  wood  and 
older  wood  was  discolored.  Other  pears,  such  as  Garber,  Tyson  and 
Surprise,  were  injured  but  little.  Since  the  freeze  the  recuperative 
ability  of  Kieffer  has  been  as  remarkable  as  was  its  susceptibility.  The 
majority  of  the  killed  spurs  have  been  replaced  by  new  growths  arising 
from  supernumerary  buds  at  the  base  of  the  old  spurs  (Fig.  13),  and 
the  spurs  whose  wood  was  blackened  in  1921  are  bearing  in  1922 
(Fig.  14). 

Since  this  variety  had  proved  so  tender  in  the  spring  of  1921,  it 
was  examined  for  injury  occurring  in  the  late  months  of  the  same 
year.  Evidence  of  injury  to  pith  near  the  tip  of  1921  wood  was  plenti- 
ful (Fig.  16),  but  there  was  no  indication  of  injury  to  fruit  buds  and 
little  or  no  indication  of  injury  to  wood.  Farther  back  on  these  same 
branches  showing  pith  injury  in  the  1921  wood,  appeared  injury  of 
another  kind.  Just  below  the  point  where  growth  was  resumed  in 
1921  both  pith  and  wood  were  injured  (Figs.  17  and  18).  Still  farther 
back,  but  yet  in  1920  wood,  the  injury  was  confined  to  the  wood.  In 
gross  appearance  there  was  a ring  of  blackened  tissue,  which  is  shown 
by  microscopic  examination  to  be  very  narrow  (Fig.  19).  Inside  the 
blackened  ring  is  a narrow  belt  of  new  wood,  one  or  two  cells  wide, 
composed  of  wood  laid  down  in  the  spring  of  1921  before  the  freeze. 
The  injury  was  confined  to  parenchymatous  tissue  and  the  wood  just 
laid  down  was  hardy  enough  to  withstand  the  freezing. 

Discoloration  of  pith  is  not  invariably  a sign  of  winter  injury, 
but,  under  the  circumstances  of  its  occurrence  in  the  material  discussed 
here,  it  may  be  taken  as  such.  The  injury  to  the  pith  in  the  1921  wood 
was  undoubtedly  an  immaturity  injury.  The  injury  to  the  pith  in  the 
1920  wood  may  have  been  due  to  the  weather  of  the  1920-1921  winter  or 


Observations  on  Winter  Injury 


15 


to  the  Easter  freeze  of  1921.  In  any  case,  it  is  clear  that  the  most 
tender  tissue  in  the  fall  of  1921  was  the  pith  while  in  the  spring  of 
the  same  year  it  was  the  wood. 

Extensive  examination  of  1920  wood  in  other  pears  and  several 
varieties  of  apple  showed  no  wood  injury  from  the  Easter  freeze  com- 
parable to  that  in  Kieffer.  In  some  cases  Ben  Davis  1920  wood  showed 
a dark  ring  (Fig.  3).  On  microscopic  examination  this  was  found  to 
be,  not  injured  tissue,  but  a false  annual  ring  caused  by  the  check  to 
growth  resulting  apparently  from  the  killing  of  the  foliage  in  the  same 
freeze. 

The  pear  trees  in  the  University  orchard  stand  within  a few  feet 
of  many  Jonathan  and  Ben  Davis  trees  and  receive  the  same  cultural 
treatment.  These  Jonathans  show  no  wood  injury  from  the  Easter 
freeze,  Ben  Davis  only  a check  to  growth,  while  Kieffer  was  severely 
affected.  In  the  fall  of  the  same  year,  however,  conditions  were  re- 
versed; Jonathan  was  injured  rather  extensively,  Ben  Davis  much  less 
and  Kieffer  least  of  all.  This  condition,  coupled  with  the  reversal  of 
the  accepted  comparative  hardiness  of  the  peach  and  the  apple,  illus- 
trates anew  the  fact  that  hardiness  is  consitutional  only  in  so  far  as 
conditions  producing  hardiness  are  constitutional.  It  is  a condition 
rather  than  a quality.  A given  fruit  is  hardy  in  a locality  as  it  reacts 
to  the  ordinary  climatic  conditions  of  that  locality  and  comparative 
hardiness  may  be  reversed  in  various  localities  according  as  the  spring 
or  the  fall  injuries  are  likely  to  prevail. 

THE  “SECOND  BLOOM” 

Following  a frost  causing  widespread  damage  to  blossoms  and 
fruit  crop  there  is  frequently  a flood  of  reports  of  crops  borne  on 
“second  bloom”  pushed  out  as  a consequence  of  the  injury  to  the  first 
crop.  The  occurrence  at  times  of  second  bloom  in  rather  considerable 
quantity  is  unquestionable.  After  a destructive  frost  it  is  naturally 
more  noticeable  than  it  would  be  in  a frostless  season  when  it  would 
come  on  about  at  the  end  of  the  first  bloom.  Its  occurrence,  however, 
is  not  necessarily  a consequence  of  frost  injury.  It  was  noted  in  great 
abundance  in  the  University  orchard  at  Columbia  in  the  spring  of  1921 
before  any  frost  had  occurred  and  in  the  spring  of  1922  when  there 
was  no  damage  from  frost.  Fig.  15,  showing  a Ben  Davis  spur  taken 
in  the  summer  of  1920,  is  significant.  Growth  for  that  year  started  at 
g.  One  apple  results  from  the  first  bloom  and  one  from  the  second. 
Clearly  in  this  case  the  destruction  of  the  first  bloom  is  not  concerned 
with  the  formation  of  the  second. 


16 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


SUMMARY 

1.  Killing  of  many  fruit  buds  in  the  apple  occurred  early  in  the 
winter  of  1921-1922  in  the  University  orchard  at  Columbia. 

2.  The  attendant  circumstances  indicate  that  this  injury  was  con- 
nected with  immaturity. 

3.  Many  plants  with  low  optimum  growing  temperatures  have, 
in  Central  Missouri,  two  distinct  growing  seasons  separated  by  a hot, 
dry  midsummer.  The  raspberry  apparently  belongs  in  this  group. 

4.  Other  plants,  with  higher  optima,  grow  rather  uniformily 
through  the  season.  This  group  includes  the  peach. 

5.  Sod  culture  may  have  the  effect  of  accentuating  the  duality 
of  the  growing  season  for  the  plants  with  low  optima.  Consequently 
its  effect  on  maturity  in  these  plants  may  be  directly  opposite  that 
recognized  in  regions  with  short  and  relatively  cool  growing  seasons. 

6.  Available  evidence  is  not  sufficient  to  indicate  how  tillage  af- 
fects maturity  in  the  apple  in  Central  Missouri. 

7.  In  the  Kieffer  pear  the  relative  hardiness  of  the  various  tis- 
sues appears  to  vary  with  the  season. 

8.  The  Kieffer  pear,  under  the  conditions  discussed  in  this  paper, 
is  more  tender  in  the  wood  in  the  spring  than  the  Jonathan  apple,  but 
more  hardy  in  fruit  buds  in  the  fall. 

9.  The  so-called  second  bloom  is  not  necessarily  the  consequence 
of  the  destruction  of  the  “first”  or  normal  bloom. 


LITERATURE  CITED 

1.  Bartlett,  G.,  Horticulturist,  1:  549.  1847. 

2.  Card,  F.  W.,  Bush  Fruits,  p.  37.,  New  York,  1917. 

3.  Chandler,  W.  H,  Proc.  Am.  Soc.  Hort.  Sci.,  12:  118.  1915. 

4.  Maynard,  S.  T.,  Agriculture  of  Massachusetts,  p.  348.  Bos- 
ton, 1884. 

5.  Whipple,  O.  B.,  Mont.  Agr.  Exp.  Sta.  Bui.  91.  1912. 


Observations  on  Winter  Injury 


17 


II. — An  Aftermath  of  Winter  Injury 

H.  A.  Cardinell 

In  the  course  of  some  continuing  demonstration  work  in  1920  and 
1921  in  a young  orchard  at  Fortescue,  Holt  County,  Missouri,  atten- 
tion was  drawn  to  the  failure  of  the  pruning  wounds  to  heal.  Where- 
ever  any  wood  was  removed,  large  or  small,  callus  formation  was 
very  slow  or  failed  altogether  (Figure  6).  Not  only  did  wounds 
which  should  have  healed  in  one  season  fail  to  cover  over,  but  the 
wood  in  the  immediate  neighborhood  seemed  dead.  Wounds  disinfected 
and  some  both  disinfected  and  painted  healed  no  better  than  those  un- 
treated. Cuts  made  below  old  wounds  revealed  much  dead  wood  in  the 
center,  surrounded  by  more  or  less  live  wood.  Cuts  through  the 
trunk  still  lower  on  the  tree,  in  many  cases  down  to  within  three  or 
four  inches  of  the  ground,  showed  the  same  condition  (Fig.  4).  For 
reasons  which  will  appear  later  in  this  paper,  it  was  possible  to  exam- 
ine 1243  trees  this  spring.  In  this  group  only  a few  cases  were  found 
of  discoloration  extending  below  the  graft  union  and  fully  50  per  cent 
of  the  trees  were  not  injured  below  three  or  four  inches  above  the 
ground  as  shown  at  k in  Fig.  4. 

Aside  from  this  failure  of  wounds  to  heal  properly  the  trees  pre- 
sented no  unusual  appearance,  except  in  some  extreme  instances.  By 
the  spring  of  1922  the  trees  most  affected,  though  many  of  these  same 
trees  were  making  20  to  40  inches  of  growth  each  year,  as  shown  in 
Fig.  5,  had  one  or  two  dead  limbs  to  the  tree.  At  the  base  of  these 
limbs  the  wood  on  the  trunks  was  practically  all  dead.  In  general, 
however,  the  trees  were  making  very  good  growth  and  on  casual  obser- 
vation the  orchard  would  have  appeared  in  excellent  condition. 

It  is  no  uncommon  occurrence  to  find  black-hearted  limbs  or 
trunks  on  trees  that  have  been  growing  and  fruiting  in  a perfectly  sat- 
isfactory manner.  This  condition  is  known  to  be  caused  by  winter 
injury,  sometimes  from  ordinary  cold  in  conjunction  with  an  immature 
condition  of  the  tree.  It  occurs  when  the  cold  is  severe  enough"  to 
kill  the  wood  but  still  not  severe  enough  to  kill  the  hardier  cambium. 
Consequently  in  the  following  spring  the  cambium  may  resume  growth 
and  surround  the  dead  area  with  a layer  of  new  tissue.  Sometimes 
these  blackened  regions  are  found  surrounded  by  Healthy  wood  show- 
ing 20  or  more  annual  rings,  indicating  that  the  injury  had  occurred 
as  many  years  previously  and  apparently  had  not  interfered  materially 


18 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


with  the  tree’s  life  or  functions.  In  itself  the  condition  is  not  serious, 
particularly  in  older  trees.  In  the  case  here  considered,  however,  there 
were  two  disturbing  circumstances:  (1)  the  failure  of  the  wounds 

to  heal,  already  mentioned,  and  (2)  the  fact  that  the  discoloration  was 
evidently  advancing  with  the  growth,  spreading  into  new  wood  (Fig. 
3,  at  B and  C).  This  made  diagnosis  of  the  cause  and  prognosis  of  the 
ultimate  effects  somewhat  uncertain. 

HISTORY  OF  THE  CASE 

The  trees  involved  stand  in  a 60-acre  block,  on  level  but  well  drained 
Missouri  River  bottom  land.  This  orchard  is  one  of  the  properties  of 
George  Hitz  & Company  of  Indianapolis  and  is  managed  by  Mr.  C.  E. 
Hitz.  In  the  spring  of  1918,  yearling  trees  to  the  number  of  2,997 
were  planted  and  in  the  fall  of  the  same  year  567  two-year-olds  were 
set.  Jonathans  were  planted  as  permanent  trees,  with  fillers  of  five 
varieties : Ben  Davis,  Delicious,  Stayman,  Grimes  and  Ingram.  All 
the  stock  used  came  from  a nursery  at  St.  Joseph,  Missouri. 

While  these  trees  were  being  cut  back  subsequent  to  planting,  Mr. 
Hitz  noticed  that  there  was  a slight  discoloration  in  the  wood.  During 
the  summer  of  the  same  year  a pathologist  from  the  United  States 
Department  of  Agriculture,  visiting  the  orchard  on  another  errand, 
examined  these  trees  and  diagnosed  the  trouble  as  winter  injury. 

DIAGNOSIS 

The  complication  already  alluded  to  and  the  resemblance  of  some 
of  the  wounds  to  cankers  of  fire  blight,  so  common  on  Jonathan  in 
Northwest  Missouri,  rather  obscured  the  case.  Specimens  of  injured 
wood  were  submitted  to  pathologists  in  various  sections  of  the  country 
and  the  possibility  of  several  other  disorders  eliminated.  In  Feb- 
ruary, 1922,  a shipment  of  injured  trees  was  sent  to  M.  B.  Waite, 
Pathologist  in  Charge,  Fruit  Disease  Investigations,  U.  S.  Department 
of  Agriculture. 

Under  date  of  February  25,  Waite  states:  “I  have  given  these 
samples  careful  study.  The  main  trouble  is  winter  injury.  It  is  com- 
plicated by  secondary  trouble  due  to  wood-rot  fungi.  These  wood-rot 
fungi  have  produced  a heart  rot  by  entering  the  frozen  injured  centers 
of  the  trunks  and  main  branches,  and  the  wood-rot  fungi  have  ex- 
tended the  injury  somewhat,  and  perhaps  complicated  and  confused 
the  primary  injury.  * * * There  are  indications  on  these  samples  that 
they  may  have  been  frozen  a second  time.  I have  often  noticed  that 


Observations  on  Winter  Injury 


19 


trees  once  injured  by  freezing  appear  slightly  more  susceptible.  Part 
of  the  two-year  wood  and  most  of  the  one-year  wood  is  sound.” 

Of  the  discoloration  in  the  one-year-old  wood  Waite  says : “This 
appears  to  be  partly  due  to  the  growth  of  the  wood-rot  fungus  from 
the  diseased  part  up  into  the  healthy  tissue.  One  of  the  samples 
shows  the  tip  of  a small  trunk  which  has  been  killed  completely  and 
shows  the  fungus  fruiting.*  Mr.  W.  H.  Diehl  of  this  bureau  has 
identified  this  fungus  as  Irpex  tulipifera  Schw” . 

Weather  records  indicate  several  possibilities  of  winter  injury 
in  the  time  since  these  trees  stood  in  the  nursery,  in  the  summer  of 

1917.  However,  inasmuch  as  injury  to  these  trees  is  known  to  have 
occurred  prior  to  planting  in  1918  and  the  secondary  injury  is  less 
certain,  interest  centers  in  the  weather  from  October,  1917,  to  March, 

1918.  The  mean  temperature  for  the  state  as  a whole  was  below  nor- 
mal during  most  of  1917  and  particularly  in  October  and  December. 
The  October  mean  temperature  was  the  lowest  on  record.  Killing 
frosts  occurred  on  October  6,  ten  days  earlier  than  the  average.  That 
fall  will  be  remembered  by  many  people  in  this  section  for  the  great 
amount  of  soft  corn. 


Table  1. — Precipitation  at  St.  Joseph,  Missouri. 
(In  Inches) 


Aug. 

Sept. 

Oct. 

Nov. 

Dec. 

1917  

5.80 

1.60 

0.80 

0.04 

0.16 

Average,  1888-1917  

4.37 

3.34 

2.62 

1.71 

0.96 

The  Weather  Bureau  records  for  St.  Joseph,  where  the  trees 
discussed  in  this  paper  were  grown,  indicate  that  the  pre- 
ceding autumn  was  rather  dryer  than  the  average.  August,  however, 
had  a rainfall  1.4  inches  above  the  average.  Whether  this  could  have 
had  any  material  effect  in  prolonging  growth  and  deferring  maturity  is 
problematical. 

Table  2. — Minimum  Temperatures  at  St.  Joseph,  Missouri. 

(In  degrees  F.) 


Oct.  Nov.  Dec.  Jan.  Feb.  Mar. 

1917-18  20  17  ^13  — 19  —73  17 

1909-1917  22  5 —10  —24  —16  -4 


•After  the  manuscript  was  prepared  many  ungrrafted  trees  had  died  and  one  type  of 
fruitincr  bodv  was  noticed  on  all.  This  was  identified  Sept.  8,  1922,  by  Diehl  as  Polystictus 
versicolor,  Fr. 


20 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


Minimum  temperatures  lower  than  any  since  1909,  when  the  St. 
Joseph  record  begins,  occurred  in  October  and  December.  Just  when 
the  injury  to  these  trees  occurred,  cannot  be  stated  definitey.  Selby 
records  extensive  damage  in  Ohio,  involving  even  complete  killing  in 
some  cases,  by  a temperature  of  18°  in  October.  Twenty  degrees  in 
October  would  seem  more  dangerous,  particularly  to  nursery  stock, 
than  — 13°  in  December,  the  other  month  of  record-making  tempera- 
ture. 

The  date  of  digging  these  trees  is  not  known.  If  they  were,  ac- 
cording to  the  prevailing  practice,  dug  in  the  fall,  the  period  of  injury 
is  fixed  without  question.  Without  this  evidence,  however,  the  prob- 
ability of  the  October  minimum  being  the  chief  factor  in  the  damage 
is  strong. 

Table  3.— Minimum  Temperatures  at  Geneva,  N.  Y. 

(In  degrees  F.) 

Oct.  Nov.  Dec.  Jan.  Feb. 


1917-18 26.0  9.0  —18.0  —10.0  —11.0 

1909-17  26.0  16.0  — 6.0  —12.0  —14.0 

1883-1909  20.5  8.0  —15.5  —18.7  —21.0 


Table  3,  compiled  from  reports  of  the  New  York  Agricultural 
Experiment  Station  at  Geneva,  which  is  a considerable  nursery  cen- 
ter and  located  in  a section  where  immaturity  is  generally  known  to  be 
the  chief  factor  in  hardiness,  is  used  here  for  comparison.  The  Octo- 
ber, 1917,  minimum  for  St.  Joseph  is  lower  than  at  Geneva  for  the 
same  year ; it  is  in  fact  a trifle  lower  than  any  in  the  long  series  of  rec- 
ords for  this  place. 

Whether  or  not  this  particular  injury  was  received  in  October,  if 
immaturity  is  likely  to  be  a factor  in  winter  injury  at  Geneva,  N.  Y., 
it  is  likely  to  be  a factor  at  St.  Joseph,  Mo.  Table  4,  giving  in  detail 
the  data  summarized  in  Tables  2 and  3,  shows  this  clearly.  In  the  nine 
years  for  which  data  are  available,  the  October  minimum  for  St.  Jo- 
seph has  been  lower  than  that  for  Geneva  in  five,  identical  in  three 
and  higher  in  one.  The  November  minimum  has  been  lower  in  five 
years,  higher  in  three  and  identical  in  one.  If  absolute  cold  at  any 
time  be  considered  the  chief  cause  of  injury,  the  Geneva  absolute 
minimum  of  — 18°F.  is  offset  by  one  of  — 24°F.  for  St.  Joseph. 

It  is  true  that  the  higher  maximum  and  mean  temperatures  of  the 
fall  months  at  St.  Joseph  may  under  certain  conditions  have  some  in- 
fluence in  hastening  maturity.  It  is  also  true,  however,  that  they  may 


Observations  on  Winter  Injury 


21 


Plate  \ . — Fig.  1 (Left)  Jonathan,  4 years  after  planting  in  the  orchard,  showing  growth  condition  and  lack  of  external 
evidence  that  would  indicate  the  condition  shown  in  Fig.  7,  a cross  section  of  the  same  trunk.  Fig.  2.  (Right)  Jonathan,  four  years 
after  planting.  Compare  this  view  with  the  cross  sections  of  the  same  tree  shown  in  Fig.  3.  Photographed  Mar.  28,  1922. 


99 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


Plate  VI. — Fig  3.  Jonathan,  four  years  from  time  of  planting,  showing 
injured  areas  and  heart  rot  rapidly  advancing  in  later  annual  rings  of  apparently 
sound  wood. 


Observations  on  Winter  Injury 


23 


Plate  VII. — big.  4.  Jonathan,  four  years  after  planting  showing  longitu- 
dinal and  cross  section  views  of  one  tree  through  trunk  and  scaffold  limbs.  In 
a large  percentage  of  the  trees  cut  off,  the  injury  terminated  in  a point  a few 
inches  above  the  ground. 


24 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


1922.  Photographed  August  14,  1922. 


Observations  on  Winter  Injury 


25 


have  some  influence  in  the  opposite  direction.  The  average  rainfall 
at  St.  Joseph  in  August,  September  and  October  is  greater  than  that 
at  Geneva ; for  these  three  months  the  figures  are,  respectively : at  St. 
Joseph,  4.37,  3.34  and  2.62  inches;  at  Geneva,  3.30,  2.42  and  2.50 
inches.  When  high  rainfall  is  combined  with  high  temperature,  growth 
is  prolonged  and  the  first  low  temperatures  are  more  likely  to  be  dam- 
aging. 


Table  4.— Minimum  Temperatures  at  St.  Joseph,  Mo.,  and  at  Geneva,  N.  Y.,  1909-1918 

Inclusive. 


Year 

October 

November 

December 

January 

February 

St.  Joseph 

Geneva 

St.  Joseph 

Geneva 

St.  Joseph 

Geneva 

St.  Joseph 

Geneva 

St.  Joseph 

Geneva 

1909-10 

27 

27 

19 

21 

—10 

1 

—13 

—8 

—5 

—3 

3910-13 

28 

26 

20 

21 

8 

—2.5 

—11 

—1 

9 

4 

1911-12 

32 

33 

5 

18 

— 4 

13 

—24 

—12 

—6 

—10 

1912-13 

31 

31 

20 

20 

8 

12 

— 4 

8 

—2 

—10 

1913-14 

22 

29 

26 

22 

11 

6 

8 

—9 

—7 

—14 

1914-15 

25 

26 

7 

16 

— 8 

—6 

—12 

—3 

13 

—10 

1915-16 

29 

29 

23 

21 

4 

4 

—19 

—3 

—4 

—8 

1916-17 

24 

29 

12 

16 

9 

4 

— 8 

1 

—16 

—8 

1917-18 

20 

(26 

17 

9 

—13 

—18 

—19 

10 

—13 

—11 

IMPORTANCE 

In  a large  number  of  cases  occurring  in  this  section  winter  injury 
of  apples  does  not  command  attention  at  the  time  of  its  occurrence. 
It  may  induce  minor  injuries,  the  consequences  of  which  are  not  re- 
vealed until  the  original  cause  is  obscured.  When  a crop  of  peach 
buds  is  killed  the  loss  is  plain,  but  when  a small  area  of  bark  is  killed 
it  receives  little  attention  until  decay  ensues  and  by  this  time  possible 
winter  injury  is  forgotten.  This  very  subtlety  of  winter  injury  makes 
difficult  any  appraisal  of  its  extent.  In  the  case  discussed  here  the  in- 
jury was  slight.  It  was  noticed  at  the  time  of  setting  the  trees,  but 
was  thought  of  no  importance.  It  did  not  affect  the  growth  of  the 
trees  and  would  have  been  forgotten  but  for  the  work  of  the  wood- 
destroying  fungi.  Many  cases  undoubtedly  occur  without  untoward 
consequence;  in  many  others  the  trees  will  grow  for  some  years  and 
when  they  begin  to  go  to  pieces  the  evidence  to  connect  the  condition 
with  a slight  winter  injury  several  years  back  will  be  scant  indeed. 

TREATMENT 

Detailed  account  of  the  treatment  given  this  orchard  following 
diagnosis  of  the  condition  will  be  published  elsewhere.  Briefly  sum- 


26 


Missouri  Agr.  Exp.  Sta.  Research  Bulletin  56 


marized,  it  consisted  in  cutting  back  to  sound  wood,  frequently  to 
within  a few  inches  of  the  ground  and  grafting  the  stubs  (Fig.  8). 
A few  trees  cut  back  without  grafting,  after  growth  had  started  and 
when  the  carbohydrate  reserve  was  low,  died.  Those  treated  earlier, 
with  grafts  inserted  in  the  crowns,  have  made  a practically  perfect 
stand  and  are  growing  vigorously.  This  procedure  has  involved  the 
sacrifice  of  the  wood  grown  in  the  four  years  these  trees  have  stood 
in  the  orchard ; but,  with  proper  attention  to  the  grafts,  it  will  ensure 
perfectly  sound  trees,  with  every  promise  of  long  life  and  productive- 
ness. 


CONCLUSIONS 

Though  winter  conditions  rarely  kill  apple  trees  outright  in  this 
section,  they  may  have  hardly  less  serious  consequences.  Evidence 
of  winter  injury  should,  therefore,  put  the  grower  on  his  guard.  If 
new  evidence  appears  every  few  years  it  may  signify  the  need  of  re- 
vision of  his  cultural  practices.  Injury  to  wood  at  any  time  justifies 
great  care  in  pruning.  If  the  cuts  can  be  made  far  enough  back  to  re- 
move all  injured  wood,  there  is  little  danger  of  infection.  If  the  re- 
moval of  all  injured  wood  is  not  practicable  there  are  two  courses  re- 
maining: (1)  careful  disinfection  and  painting  of  all  wounds,  (2) 
omission  of  pruning  altogether  till  the  cuts  can  be  made  in  sound  tissue 
growing  subsequent  to  the  freeze.  The  practicability  of  these  methods 
will  be  discussed  elsewhere.  However,  one  guiding  principle  may  be 
stated:  injured  wood  should  not  be  exposed.  Sealed  within  living 
wood,  it  is  harmless ; exposed  it  is  a source  of  constant  danger. 


UNIVERSITY  OF  MISSOURI  COLLEGE  OF  AGRICULTURE 
AGRICULTURAL  EXPERIMENT  STATION 
RESEARCH  BULLETIN  56 


Observations  on  Winter  Injury 


I — Early  and  Late  Winter  Injury 
II — An  Aftermath  of  Winter  Injury 


COLUMBIA.  MISSOURI 
NOVEMBER,  1922 


46-M  Ik].22UETIN  COLOmIia.  MO. 


3 0112  019681935 


